UvA-DARE (Digital Academic Repository)

Transcription

1 UvA-DARE (Digital Academic Repository) Pollen-based biome reconstructions for Latin America at 0, 6000 and radiocarbon years Marchant, R.A.; Harrison, S.P.; Hooghiemstra, H.; Markgraf, V.; van Boxel, J.H.; Ager, T.; Almeida, L.; Anderson, R.; Baied, C.; Behling, H.; Berrio Mogollon, J.C.; Burbridge, R.; Björck, S.; Byrne, R.; Bush, M.B.; Cleef, A.M.; Duivenvoorden, J.F.; Flenley, J.R.; de Oliveira, P.; van Geel, B.; Graf, K.J.; Gosling, W.D.; Harbele, S.; van der Hammen, T.; Hansen, B.C.S.; Horn, S.P.; Islebe, G.A.; Kuhry, P.; Ledru, M-P.; Mayle, F.E.; Leyden, B.W.; Lozano-García, S.; Melief, A.B.M.; Moreno, P.; Moar, N.T.; Prieto, A.; van Reenen, G.B.A.; Salgado-Labouriau, M.L.; Schäbitz, F.; Schreve-Brinkman, E.J.; Wille, M. Published in: Climate of the Past Discussions DOI:.194/cpd Link to publication Citation for published version (APA): Marchant, R., Harrison, S. P., Hooghiemstra, H., Markgraf, V., van Boxel, J. H., Ager, T.,... Wille, M. (09). Pollen-based biome reconstructions for Latin America at 0, 6000 and radiocarbon years. Climate of the Past Discussions, (1), DOI:.194/cpd General rights It is not permitted to download or to forward/distribute the text or part of it without the consent of the author(s) and/or copyright holder(s), other than for strictly personal, individual use, unless the work is under an open content license (like Creative Commons). Disclaimer/Complaints regulations If you believe that digital publication of certain material infringes any of your rights or (privacy) interests, please let the Library know, stating your reasons. In case of a legitimate complaint, the Library will make the material inaccessible and/or remove it from the website. Please Ask the Library: or a letter to: Library of the University of Amsterdam, Secretariat, Singel 4, 12 WP Amsterdam, The Netherlands. You will be contacted as soon as possible. UvA-DARE is a service provided by the library of the University of Amsterdam ( Download date: 02 Jul 18

2 Clim. Past Discuss.,, , 09 Author(s) 09. This work is distributed under the Creative Commons Attribution 3.0 License. Climate of the Past Discussions Climate of the Past Discussions is the access reviewed discussion forum of Climate of the Past Pollen-based biome reconstructions for Latin America at 0, 6000 and radiocarbon years R. Marchant 1, S. P. Harrison 2, H. Hooghiemstra 3, V. Markgraf 4, J. H. van Boxel 3, T. Ager, L. Almeida 6, R. Anderson 7, C. Baied 8, H. Behling 9, J. C. Berrio, R. Burbridge 11, S. Björck 12, R. Byrne 13, M. B. Bush 14, A. M. Cleef 3, J. F. Duivenvoorden 3, J. R. Flenley, P. De Oliveira 16, B. van Geel 3, K. J. Graf 17, W. D. Gosling 18, S. Harbele 19, T. van der Hammen 3,, B. C. S. Hansen 21, S. P. Horn 22, G. A. Islebe 23, P. Kuhry 24, M.-P. Ledru, F. E. Mayle 26, B. W. Leyden 32, S. Lozano-García 27, A. B. M. Melief 3, P. Moreno 28, N. T. Moar 29, A. Prieto 30, G. B. van Reenen 3, M. L. Salgado-Labouriau 31, F. Schäbitz 33, E. J. Schreve-Brinkman 3, and M. Wille 33 1 The York Institute for Tropical Ecosystem Dynamics (KITE), Environment Department, University of York, York, Heslington, YO DD, UK 2 Bristol Research Initiative for the Dynamic Global Environment (BRIDGE), School of Geographical Sciences, University Road, University of Bristol, Bristol BS8 1SS, UK 3 Institute for Biodiversity and Ecosystem Dynamics (IBED), Faculty of Science, University of Amsterdam, Postbus 94062, 90 GB Amsterdam, The Netherlands INSTAAR, University of Colorado, Boulder, Colorado 80309, USA USGS, National Centre, MS 970, Reston, Virginia 292, USA 6 Laboratorio Biogeografía, Facultad de Ciencias, Universidad Nacional Autónoma de México, Aptdo Postal , 04 México D.F., México 7 Department of Geography, University of Montana, Missoula, Montana , USA 8 Environmental Studies Program, University of Montana, Missoula Montana 9812, USA 9 Georg-August-Universität, Albrecht-von-Haller-Institut für Pflanzenwissenschaften, Abteilung Palynologie und Klimadynamik, Untere Karspüle 2, Göttingen, Germany Department of Geography, University Road, University of Leicester, LE1 7RH, UK 11 c/o. Geography Building, Drummond Street, Edinburgh, EH8 9XP, UK 12 Geological Institute, Univ. of Copenhagen, Øster Volgade, 130 Copenhagen, Denmark 13 Department of Geography, University of California, Berkeley, California , USA 14 Department of Biological Sciences, Florida Institute of Technology, 0 W. University Boulevard, Melbourne, Florida 3290, USA Department of Geography, Massey University, Palmerston, New Zealand 16 Instituto de Geociencias-DPE, Universidade de São Paulo, Caixa Postal 11348, São Paulo, SP , Brazil 17 Geographisches Institut der Universität, Winterthürerstrae 190, 807 Zürich, Switzerland 18 Department of Earth and Environmental Sciences, CEPSA, The Open University, Walton Hall, Milton Keynes, MK7 6AA, UK 19 Dept. of Geography and Environmental Science, Monash Univ., Clayton, Victoria, Australia Fundación Tropenbos Colombia, Carrera 21 #39-3, Santafe de Bogotá, Colombia 21 Limnological Research Centre, University of Minnesota, 2 Pillsbury Hall, 3 Pillsbury Drive, Minneapolis, Minneapolis , USA 22 Department of Geography, University of Tennessee, 408 G&G Building, Knoxville, Tennessee , USA 370

3 23 El Colegio de la Frontera Sur. ECOSUR-Chetumal, Apartado Postal 424, Chetumal, Quintana Roo, CP 77000, México 24 Department of Physical Geography and Quaternary Geology, Stockholm University, 691 Stockholm, Sweden Equipe Paléoenvironnements, Institut des Sciences de l Evolution Institut de Recherche pour le Developement, Montpellier, France 26 Geography Building, Drummond Street, Edinburgh, EH8 9XP, UK 27 Universidad Nacional Autónoma de México, Instituto de Geología, Aptdo Postal , 04 México D.F., México 28 Facultad de Ciencias, Universidad de Chile, Casilla 63, Santiago, Chile 29 Botany Division, D.S.I.R., Private Bag, Christchurch, New Zealand 30 Laboratorio de Palinologia, National Universidad Mar del Plata, Departmento de Biologia, Funes 30, 7600 Mar del Plata, Argentina 31 Instituto de Geociencias, Fundação Universidade do Brazilia, Campus Universitario, Asa Norte, , DF Brazilia, Brazil 32 Department of Geology, University of South Florida, Tampa, FL 336, USA 33 Seminar für Geographie, Universität zu Köln, Gronewaldstrasse 2, 0931 Köln, Germany Received: 13 October 08 Accepted: 21 October 08 Published: February 09 Correspondence to: R. Marchant (rm24@york.ac.uk) Published by Copernicus Publications on behalf of the European Geosciences Union. 371 Abstract The biomisation method is used to reconstruct Latin American vegetation at 6000±00 and ±00 radiocarbon years before present ( 14 C yr BP) from pollen data. Tests using modern pollen data from 381 samples derived from 287 locations broadly reproduce potential natural vegetation. The strong temperature gradient associated with the Andes is recorded by a transition from high altitude cool grass/shrubland and cool mixed forest to mid-altitude cool temperate rain forest, to tropical dry, seasonal and rain forest at low altitudes. Reconstructed biomes from a number of sites do not match the potential vegetation due to local factors such as human impact, methodological artefacts and mechanisms of pollen representivity of the parent vegetation. At 6000±00 14 C yr BP samples are analysed from 127 sites. Differences between the modern and the 6000±00 14 C yr BP reconstruction are comparatively small. Patterns of change relative to the modern reconstruction are mainly to biomes characteristic of drier climate in the north of the region with a slight more mesic shift in the south. Cool temperate rain forest remains dominant in western South America. In northwestern South America a number of sites record transitions from tropical seasonal forest to tropical dry forest and tropical rain forest to tropical seasonal forest. Sites in Central America also show a change in biome assignment to more mesic vegetation, indicative of greater plant available moisture, e.g. on the Yucatán peninsula sites record warm evergreen forest, replacing tropical dry forest and warm mixed forest presently recorded. At ±00 14 C yr BP 61 samples from 34 sites record vegetation that reflects a generally cool and dry environment. Cool grass/shrubland prevalent in southeast Brazil, Amazonian sites record tropical dry forest, warm temperate rain forest and tropical seasonal forest. Southernmost South America is dominated by cool grass/shrubland, a single site retains cool temperate rain forest indicating that forest was present at some locations at the LGM. Some sites in Central México and lowland Colombia remain unchanged in their biome assignments, although the affinities that 372

4 these sites have to different biomes do change between ±00 14 C yr BP and present. The unresponsive nature of these sites results from their location and the impact of local edaphic influence. 1 Introduction Biomisation is an objective method to reconstruct broad vegetation types based on the assignment of pollen taxa to one or more plant functional types (PFTs) (Prentice et al., 1996a). The method is based on the assumption that a pollen spectrum will have different degrees of affinity to different biomes that can be quantified by a simple algorithm. Biome reconstructions from pollen data at 6000±00 14 C yr BP and the last glacial maximum (LGM) at 00± C yr BP have been produced for most regions of the world under the auspices of the BIOME 6000 project (Prentice et al., 1998, 00). The validity of the method in reconstructing biomes at different time intervals has been demonstrated for Africa (Jolly et al., 1998a; Elenga et al., 00), Australia (Pickett et al., 04) Beringia (Bigelow et al., 03; Edwards et al., 00), China (Yu et al., 1998, 01), Eastern North America (Williams et al., 00), Eurasia (Tarasov et al., 1998a), Europe (Prentice et al., 1996a, b; Tarasov et al., 1998a, b; Elenga et al., 00), Japan (Takahara et al., 01) and Western North America (Thompson and Anderson, 00). Results from Latin America, presented here, represent the last geographically large area to undergo this process. Within Latin America the biomisation method has been previously applied to Colombian pollen data at a range of spatial and temporal scales; from the middle Holocene (Marchant et al., 01a), the LGM (Marchant et al., 02a), to investigate modern-pollen vegetation relationships (Marchant et al., 01b), impact of human societies on vegetation (Marchant et al., 04a) and as a basis for comparisons with output from a vegetation model run under different climatic and environmental scenarios (Marchant et al., 04b, 06). In addition to these spatial investigations, the method has been applied down-core down to a year pollen record from the high plain of Bogotá (Marchant et al., 02b). As Colombia is biogeograph- 373 ically complex, encompasses high altitude, temperate and tropical floras reflecting a range of environmental space including transitions from hyper-humid to semi-arid climates, these analyses provided a suitable test-bed for the wider geographical focus presented here. In addition to reconstructing vegetation patterns, and investigating factors that can explain observed changes, data on past biomes contributes to testing of climate and vegetation models (Prentice et al., 1992; Haxeltine and Prentice, 1996; Peng et al., 1998; Marchant et al., 06; Braconnot et al., 07). Vegetation models can be used to portray output from Global Circulation Models (GCMs) as maps of potential vegetation (Claussen and Esh, 1994; Foley et al., 1996; Prentice et al., 1996b; Williams et al., 1998) that can be used in the development of models that couple biosphere, atmosphere and oceanic components (Braconnot et al., 07; Claussen, 1994; Harrison et al., 03; Texier et al., 1997) and testing of biogeochemical dynamics (Peng et al., 1998). There has been growing interest has focused how atmosphere-biosphere interactions have operated under the changing environmental conditions since the LGM, particularly in trying to understand the response of ecosystems to different types of environmental forcing (Jolly and Haxeltine, 1997). Transformed pollen data can further be used in conjunction with other data types, such as on lake status (Jolly et al., 1998b) and archaeological evidence (Piperno et al., 1990, 1991a, b), to better understand the causal factors driving vegetation change over the recent geological past. 1.1 Latin American region Latin America comprises the area from 3 N to 6 S, and from 3 W to 1 W extending from México to islands off southernmost South America from eastern Brazil to the Galapagos Islands. Latin America is characterised by strong environmental gradients associated with 0 of latitude, approximately 7000 m of altitude and the transition from oceanic- to continentally-dominated climate systems (Fig. 1). Physiographically, Latin America is characterised by stable cratons associated with the interior and areas of active mountain building, particularly associated with the Andes. This environmental 374

5 variability is reflected by an incredibly diverse biogeography, ranging from the highly diverse rain forest of the Chocó Pacific (Colombia) to the cold deserts of the high Andes, from the hot semi-desert areas of México to the cold moorlands of Tierre del Fuego (Fig. 2). Descending an altitudinal gradient there is a transition from páramo (cool grass/shrubland) to high Andean forests (cool mixed and cool temperate rain forests) and lower Andean forest (warm evergreen forest) (Fig. 3). Complicating this potential vegetation distribution is the factor of human impact with the majority of the vegetation in Latin America being impacted on by the vegetation (Ellis and Ramankutty, 08). The timing of early human settlement in Latin America is a contentious subject, although it seems from the early Holocene there was considerable cultural diversity and adaptation to a series of different environments (Gnécco, 1999). Human-induced impact has had a direct influence on vegetation composition and distribution through land-use practices and the introduction of alien taxa and cultivars to the Latin American flora. For example, in excess of 0 plants were under cultivation prior to the European conquests in the th century (Piperno et al., 00) Latin America climate Cerverny (1998), Eidt (1968) and Metcalfe et al. (00) have reviewed Latin American climate. Given the broad geographical scope, Latin America is characterised by a variety of climates that relates to its global position, shape of the landmass, location and height of the Andes, offshore currents, general hemisphere air flow and proximity of large water bodies (Fig. 1). Four dominant circulation regimes influence Latin America: the Inter-Tropical Convergence Zone (ITCZ), the prevailing westerlies, the semi-permanent high pressure cells located over the South Pacific and South Atlantic Oceans and the trade winds. Perhaps most dominant is the annual oscillation of the meteorological equator (ITCZ), this migrating some latitude about the equator (Fig. 1). The ITCZ reaches its northernmost location in June, this bringing high rainfall for northern South America and the Caribbean, with January and February recording the dry season (Cerveny, 1998). However, due to the influence of the westerlies from 37 the Pacific, and the sharply rising topography of the Andes, the ITCZ has a sinusoidal profile over northwestern South America (Fig. 1). In southern South America the prevailing westerlies south of 40 S are particularly important in controlling the moisture regime. The topographic barrier of the Andes contributes to the creation of two large semi-present anticyclones, one over the South Pacific and one over the South Atlantic, the southeast trade winds associated with this latter system brings abundant moisture to the Amazon Basin (Cerveny, 1998). Due to the large size of South America, and the highland ranges that fringe much of the continent there is often a rapid transition from relatively moist coastal areas to a dry interior reflecting the transition from oceanic- to continental-dominated climate systems. For example, due to the proximal location of the Pacific-based moisture source and steeply rising ground, precipitation is highest (> 000 mm yr 1 ) in the Chocó Pacific region. Exceptions to this scenario are areas located between the anticyclones, e.g. the Peruvian coast, where relatively arid conditions prevail. One of the main environmental gradients in Latin America is associated with the Andes. The Andes are characterised by a diurnal climate (Kuhry, 1988); at a given location differences in monthly temperature are small (<3 C) although daily fluctuations may be large ( C), especially during the dry seasons. Climatic changes with altitude can be summarised as a lapse rate (Barry and Chorley, 1990). Applying a lapse rate of 6.6 C 00 m 1 (Van der Hammen and González, 196; Wille et al., 01), this altitudinal rise equates to a temperature change of more than 30 C. Also associated with the Andes are steep gradients of moisture availability. Rainfall is high on the eastern slopes of the Andes; the concave nature acting as a receptacle for moisture transferred by the southeast trade winds from the Atlantic Ocean, in part receiving moisture generated by the Amazonian forest (Fjeldså, 1993). Low rainfall is recorded within rain shadow areas, such as on the lower slopes of the Magdalena Valley and the inter-andean plains (Kuhry, 1988). These climate gradients result in rapid transitions from mesic to xeric vegetation types, e.g., cool high-altitude grasslands change to temperate forests at mid-altitudes and diverse tropical rain forests within a few kilometres (Fig. 3). In south- 376

6 ern part of Latin America rainfall is largely controlled by the persistence and strength of the westerly winds (Gilli et al., 0). In recent years there has been increased interest in large-scale temperature-driven surface pressure oscillations in the Pacific Ocean termed the Southern Oscillation, and its assimilated oceanic aspects, El Niño and its antithesis La Niña (Cerveny, 1998). The climate of the tropical Pacific basin, extending from the western Americas across to Australia, New Zealand, and northeast Asia, oscillates at irregular time intervals (3 to 7 years) between an El Niño phase, with warm tropical waters upwelling off Pacific coastal South America, and a La Niña phase, with cold tropical waters dominating. ENSO events are the largest coupled ocean-atmosphere phenomena resulting in climatic variability on inter-annual time scales (Godínez-Domínguez et al., 00). As climates, particularly rainfall patterns, are driven by temperature differences between land and ocean, the influence of changing oceanic sea surface temperatures (SST) on coastal South American environment can be dominant, and have a strong influence elsewhere (Marchant and Hoohiemstra, 04). El Niño events primarily result in increased precipitation along the Pacific coastal regions, decreased precipitation within lowland tropical moist forests of Central America (Cerveny, 1998) with increased precipitation in northern Central America (Metcalfe et al., 00) Latin America vegetation For the purpose of this investigation the potential vegetation composition and distribution Latin America is classified at a coarse resolution with twelve biomes being identified (Fig. 2) that summarise the 7 categories mapped by Hück (1960) and 4 by Schmithüsen (1976). The vegetation composition and distribution generally reflects the main climatic and topographic gradients described above. However, a series of caveats to this must be stressed. Firstly, the actual and potential vegetation can be quite different, the former reflecting a long history of human interaction that has been particularly pronounced since the colonial period but has been influencing the vegetation for at least the last 000 years (Marchant et al., 04). In numerous areas this 377 interaction has completely transformed the potential vegetation to an agricultural landscape. Another factor complicating the relationship between climate and vegetation is the locally strong edaphic influence by substrate, topography or geographic character (Fig. 3). The strength of this influence is characterised by areas of tropical dry forest that forms on free-draining sandstones, e.g. the Llanos Orientales (Colombia); these are located in areas where the climate regime would support tropical seasonal forest, or even tropical rain forest. The vegetation at such locations is relatively insensitive to climate changes as these must be of a greater magnitude than the influence imparted by the edaphic factor. Broad types of vegetation with similar composition and distribution (biomes) result from a combination of plant functional types (PFTs). PFTs and biomes, which lie at the heart of the biomisation technique, allow the high floristic diversity of the Latin America pollen flora to link with the relatively coarse vegetation classification (Fig. 4). PFTs group together species that have common character (Prentice et al., 1992). This grouping is based on common life form and phenology, combined with the geographic distribution that is in part determined by climate (Woodward, 1987). An indication of the bioclimatic range of each PFT and plant physiological adaptation, to the given environmental condition, is presented in Table 2. The range of biomes identified within the Latin America, floristic description, main location and equivalent floristic units is portrayed in Table 3. The cool grass/shrubland biome incorporates a relatively wide range of vegetation dominated by grasses, heath, cool temperate sclerophyll shrubs and cushion plants (Fig. 3). This biome is present in southern South America and at high altitudes along the Andes. In addition to the cool grassland, a warm grassland (steppe) is identified. Steppe is found predominately under the warm, dry climates of southeast and northeast Brazil, northwestern Argentina and coastal northern South America. Warm temperate rain forest represents a mix of warm conifers such as Araucaria, Andean and Atlantic rain forest elements, whereas cool temperate rain forest contains cool conifers, such as Fitzroya, Andean and Valdivian rain forest elements. Dry forests are extensive in Latin America, specifically associated with areas located 378

7 between the two semi-permanent anticyclones and influenced by the high seasonality of rainfall imposed by the annual migration of the ITCZ. For our classification we characterise the diverse dry vegetation formations (Fig. 4) as the tropical dry forest and xerophytic trees and shrub biomes. Xerophytic trees and shrubs is widespread in the interior of South America, along the southwestern Pacific coast and northeast Brazil where it grades into steppe, additionally, there are patches in Colombia, on the Yucatán peninsula and in México (Fig. 2). Tropical dry forest is predominantly recorded in two main swaths either side of the Amazon basin, with an extension through Central America. The tropical seasonal forest biome is predominantly recorded to the north of Amazonia where it is interspersed with patches of dry forest; this reflecting a strong edaphic influence. A large area of tropical seasonal forest is recorded away from the hyper-humid area of Brazil along the Atlantic coast. The tropical rain forest biome is present in three main areas: Amazonia, linear strips along the Atlantic coast and northeast South America extending into Central America. Forest associated with highland areas is divided into three biomes: warm evergreen forest, cool temperate rain forest and cool mixed forest (Fig. 4). Warm evergreen forest is most extensive along the lowland Andes, adjacent to the tropical rain forest. Cool mixed forest has a more restricted distribution, occupying a highland position until temperature becomes limiting for a number of taxa. Warm mixed forest is characterised by a mix of Pinus and Quercus species and is mainly restricted to Central America. The desert biome is restricted to coastal Peru, due to the Pacific Ocean anticyclone, this area receives very little moisture, except when the area is subjected to El Niño events. 2 Methods 2.1 Data sources Over the past five decades palynologists have collected numerous pollen-based records from lakes and bogs (Table 1) that have been used to unravel past vegeta- 379 tion changes in Latin America with ecosystem reconstructions now existing from all major vegetation types over the Late Quaternary period. The Latin American Pollen Database (LAPD) ( is an online resource used to collate these data and facilitated the systematic interoperation presented here; indeed the majority of the pollen data used here are available through the LAPD. Additional data were obtained from palynologists working in Latin America; all active palynolgists being given the opportunity to contribute data not currently lodged in the LAPD. Indeed, data from a number of sites in Argentina, Brazil, Costa Rica, México and Panama were made available specifically for this work. The majority of data from Colombia were prepared for this analysis directly from the original count sheets and are in preparation for uploading to the LAPD. The majority of the data used in our analysis are complete raw pollen counts, this permitted all pollen taxa recorded by the original analyst to be allocated to PFTs and allowed the integrity of the data to be maintained throughout the analysis. Application of raw pollen data in other regions has been shown to help in differentiating between biomes (Tarasov et al., 1998b). However, numerous pollen records are either not submitted to the LAPD, or, were not made available for this analysis. Rather than omitting these data, the pollen counts were digitised from published pollen diagrams (Table 1): digitising such data provides a spatially more complete reconstruction than available from presently archived data. The process of digitisation involved either back calculation of the pollen counts if information on the pollen sum was present. If the pollen sum was not available the pollen percentage diagram was used as a count of 0 and values for the pollen taxa were abstracted at the time intervals used for our analysis. This scenario of combining data from different formats comes with a number of caveats that can have bearing on the results, and their interpretation (Marchant and Hooghiemstra, 01). Firstly, the sub-set of pollen taxa in a count used to construct published pollen diagrams, and pollen sums that comprise it, often result from the bias of individual researchers, particularly on what are the reliable indicator taxa for a particular area and range of different vegetation types under investigation. This issue is particularly crucial 380

8 in Latin America where the large numbers of pollen taxa encountered in the original counting are rarely depicted on published pollen diagrams. Furthermore, the level of identification achieved within pollen analysis, to a generic or family level, commonly comprises species that can be found in a range of different vegetation types, ecologies and growth forms (Marchant et al., 02c). The majority of the samples for the biomisation presented here are derived from sites close to the Andean spine. Primarily, this concentration reflects the sensitive response of the vegetation to climate change on the steep altitudinal gradients (Marchant et al., 01b); the area forming an ideal location for palaeoecological research. Additionally, the comparative lack of data from the lowlands is fuelled by problems of access, suitable sites and strong river dynamics that commonly result in sedimentary hiatuses (Ledru, 1998). This spatial bias of the location did not reduce the number of biomes we were able to reconstruct, because of the steep environmental gradients associated with 7000 m of altitudinal change found along the Andes (Fig. 4). Uncalibrated radiocarbon dates available from the original stratigraphic analysis were used to select samples representing the time period used here; these were. On a site-by-site basis, a linear age-depth model was applied to the pollen data. The validity of this model was assessed at each site taking into account sedimentary hiatuses and dating problems such as age reversals and dates with large standard errors; a summary of this dating control is provided in Table 6 following the COHMAP scheme (Webb, 199; Yu and Harrison, 1999). Multiple samples ( 3) were selected when more than one sample fell within the age range allowed for each time period. These data were compiled, to produce a site vs. taxa matrix that was then checked to standardise nomenclature, e.g., the combined file contained many synonyms such as Gramineae and Poaceae, and Mysine and Rapanea. Synonymous taxa were combined using the nomenclature of Kewensis (1997) and the International Plant Names Index (IPNI) (1999). Aquatic and non-arboresent fern taxa were removed from the matrix as they commonly reflect local hydrological conditions rather than local climate envelope. Marker additions and exotic spikes such as Lycopodium were also removed. 381 A total of 381 samples from 287 locations derived from core tops (<00 14 C yr BP), surface samples, pollen traps and moss polsters comprise the modern data set (Table 1). For the time period 6000±00 14 C yr BP, samples derived from 127 pollen records comprise the data set (Table 1). For the time period ±00 14 C yr BP, 61 samples derived from 34 pollen records comprise the data set (Table 1). The data sets to undergo analysis comprised pollen taxa for the modern calibration, 493 for the C yr BP reconstruction and 232 for the C yr BP reconstruction. The taxonomic diversity of the Neotropical phytogeographical realm can be demonstrated by taking the modern biomisation as an example: the number of pollen taxa for the production of our biomes is greater than Africa (364) (Jolly et al., 1998a), Europe (41) (Prentice et al., 1996b), Russia and Mongolia (98) (Tarasov et al., 1998a) and China (68) (Yu et al., 1998). 2.2 Biomisation Prentice et al. (1996a) and Prentice and Webb (1998) have documented the steps involved in the biomisation technique. First, a conceptual framework for biomes and PFTs in Latin American vegetation was developed by investigating the relationship between potential biomes and three environmental gradients. The environmental gradients considered were moisture availability (α: Priestley-Taylor coefficient of plant available moisture), temperature (MTCO: mean temperature of the coldest month) and seasonal warmth (GDD: growing degree-days). To enable a definition of the biomes to be based on bioclimatic data, rather than qualitative assessment, the climate space encompassed by Latin America was plotted against climate data set of Leemans and Cramer (1991) with relationship between biomes at individual site locations and macroscale climate changes (α and MTCO) investigated in two-dimensional space (Fig. ). The twelve biomes identified within Latin America (Table 2) are designed to incorporate the range of major vegetation types and ensure consistency with previous areas to undergo the process within the BIOME 6000 community. 382

9 Similarly to the biomes, but on a finer ecological resolution, the spatial distribution of PFTs is determined by environmental controls on plant growth form and ecological tolerance (Woodward, 1987). In Latin America the dominant environmental gradients are temperature, primarily associated with altitude, moisture availability and seasonality. PFT definitions were modified from the classification originally developed for the BIOME 1 model (Prentice et al., 1992, 1996a, b) taking into account schemes developed for other regions, particularly those that abut the Latin American region or contain similar floristic elements (Jolly et al., 1998a; Pickett, et al., 04; Takahara et al., 01 Elenga et al., 00; Thompson and Anderson, 00; Yu et al., 00). Five main groups of PFT were distinguished: these containing tropical (non-frost tolerant), coniferous (needle-leaved), temperate (frost tolerant), xerophytic (drought tolerant), and frost and drought tolerant taxa (Fig. 6). This latter group is present in cold dry conditions of southern South America and the high Andes. A sixth miscellaneous group represents various life forms with restricted diagnostic value. The Latin American flora was divided into PFTs (Table 3). The PFTs, although being ecological distinct, can be multiply assigned to the biomes (Table 4). The classification is based on the original scheme devised for the Biome 3 vegetation model (Prentice et al., 1992) and modification through regional applications to pollen data. Where possible the scheme devised for Latin America conforms to existing classification and definitions. However, some of the specific vegetation types in Latin America were not adequately covered by the existing range so two new PFTs (heath and cushion plants) were added. To aid in the separation of the African forest/savanna boundary, Jolly et al. (1998a) subdivided the tropical raingreen trees PFT (Tr) into three groups. In the case of Latin America, it was decided that the overlap (taxa being multiply assigned to the PFTs) between the PFTs would be too great, and the distinction somewhat minimal. Furthermore, the tropical xerophytic trees and shrubs PFT encompass many taxa that would be assigned to the driest tropical raingreen category. Therefore, the Tr PFT was subdivided into wet (Tr 1 ) and dry (Tr 2 ) tropical raingreen trees. The cornerstone of research concerned with the composition and distribution of Latin 383 American vegetation, be it in a contemporary time frame, or one that aims to work in the past, is a good understanding of the ecology and distribution of the taxa concerned. The Latin American pollen taxa were assigned to one or more PFTs depending on the modern ecological range of the most important (i.e. most abundant) taxa responsible for producing the pollen identifiable within the modern data set. These assignments were made following reference to the known biology of plants from several floras (Rzedowski, 1983; Schofield, 1984; Wingenroth and Suarez, 1984; Kahn and de Granville, 1992; Gentry, 1993; Maberly, 1993; Seibert, 1996), botanical and palynological studies (Beard, 19; van der Hammen, 1963, 1972; Wijmstra and van der Hammen, 1966; Eiten, 1972; Cleef and Hooghiemstra, 1984; Hooghiemstra and Cleef, 1984; Pires and Prance, 198; Prance, 198; Cuatrecasas and Barreto, 1988; Brown and Lugo, 1990; Bush, 1991; Dov Par, 1992; Kappelle, 1993; Duivenvoorden and Cleef, 1994; Witte, 1994; Armesto et al., 199; Harley, 199; Kappelle, 199; Kershaw and McGlone, 199; Veblen, et al., 199; Colinvaux, 1996; Grabherr, 1997; Hooghiemstra and van der Hammen, 1998) and personal communication with modern ecologists and palaeoecologists. Much of this information has been collated into a dictionary on the distribution and ecology of parent taxa of pollen lodged within the Latin American Pollen Database (Marchant et al., 02c). The resultant taxon vs. PFT assignments are presented in Table. Due to the high intra-generic diversity, and also the wide range of ecology s exhibited by the parent taxa present within some genera, a number of taxa were multiply assigned to a number of PFTs; where possible, pollen taxa were assigned to the PFTs within which the parent taxa are most common. Thus, the identified PFTs from Latin America are described by the suite of pollen taxa assigned to them, in turn the biomes are distinguished by the suite of constituent PFTs. A number of pollen taxa belong to more than one PFT, and, as is the case with the potential vegetation, most PFTs contribute to more than one biome. Two problems can arise here for our analysis that can be circumvented by manipulation of the input matrices and output biome affinity scores. First, pollen samples can have equal maximum affinity with more than one biome; this commonly occurs when the 384

10 PFTs characteristic of one biome are a subset of another biome. Assigning the biomes so that subsets always come first in the analysis solves the problem. A second problem arises where multiple samples encompass the age boundaries. Multiple samples from a single site may have maximum affinity to a number of different biomes; the chance of this is high when the score of the best biome is close to that of the next best. In such cases, the majority biome is mapped. For example, site A contains eight samples within the time frame of 6000±00 14 C yr BP, five samples have the greatest affinity to biome 1, two samples to biome 2 and one sample to biome 3. The result is that biome 1 is mapped for site A at 6000±00 14 C yr BP. Biomes were reconstructed from pollen data at sites with surface sample, trap and radiocarbon-dated core-top data. The results were used to produce a modern pollenderived biome dot map (Fig. 7); for each site a colour dot records the reconstructed biome with the highest affinity score. These were compared site by site, with the potential modern vegetation distribution (Fig. 2). The biomisation procedure was applied to the fossil datasets without modification. Results for all sites and periods are provided in Table 6 which allows a site-by-site comparison through time and a comparison between the modern reconstruction and potential vegetation. 3 Results 3.1 Modern pollen vs. potential biome reconstruction Visual comparison shows that the biomes reconstructed from modern pollen data (Fig. 7) accurately reflect the broad features in the potential vegetation map (Fig. 2). In particular the modern reconstruction correctly reproduces the transition from relatively mesic vegetation types, around the coastal areas of South America, to the more xeric biomes towards the interior. For example, in eastern Argentina there is a transition from steppe to xerophytic woods and scrub. Warm temperate rain forest is an important biome in the southern and southeastern Brazilian highlands, with tropical dry forest 38 being reconstructed towards the interior. Notable from this region is the large number of different biomes being reconstructed in a relatively small area. In part this reflects the variability of potential vegetation, not portrayed in our relatively coarse resolution vegetation map (Fig. 2). For example, a site recording tropical rain forest reflects the sites lowland position where it is characterised by moist gallery forest with a number of typical rain forest taxa present. Steppe is correctly reconstructed from the grasslands of south-eastern Argentina and dry forest in central Argentina, mirroring the transition to drought-deciduous thorn forests of central Argentina (Schmithüsen, 1976). Steppe is assigned farther west at approximately 00 m in the Andes, southernmost South America and northeast Brazil. The vegetation of southern South America is dominated by cool temperate rain forest. The failure of the analysis to pick up the transition from cool temperate rain forest to cool grass/shrubland as one progresses east along Tierre del Fuego stems from the pollen spectra having a relatively large amount of Nothofagus pollen. Moving northwards from southern South America there is a transition to cool mixed forest, cool grass/shrubland and steppe; these latter assignments are particularly associated with eastern flanks of the Cordillera de los Andes. The concentration of sites along the Andes results in a wide range of reconstructed biomes being geographically adjacent to each other when mapped in two-dimensional space (Fig. 7). This phenomenon is most apparent in the northern Andes where the altitudinal, and therefore climatic gradients are at their steepest. Despite these rapid environmental changes, the biome assignments reflect the changing vegetation patterns. There is a clear altitudinal transition: low altitudes (<300 m) being mainly assigned to the tropical rain forest, tropical dry forest, tropical seasonal forest and steppe. Sites located at mid altitudes are described by a number of different biomes including tropical seasonal forest, warm mixed forest, cool mixed forest and cool temperate rain forest. Within this wide range of warm temperate rain forest and tropical seasonal forest are commonly assigned at lower elevations (Fig. 7, Table 6). Many of the sites at high altitude have a high affinity to the cool grass/shrubland biome. The line of the Andes can be easily seen by the cool grass/shrubland biome assignments, these being commonly 386

11 recorded at sites above 3800 m. The warm temperate rain forest is assigned at lower elevations and is analogous to Andean forest, being dominated by Podocarpus, Quercus and Weinmannia and comprises a different forest type to that assigned in southern and southeast Brazil or southern South America. A mixture of biomes presently characterises the Amazon Basin with only four sites recording the tropical rain forest biome; two of these are in coastal locations. Tropical seasonal forest is recorded in four locations; this representing a slightly drier type of forest than tropical rain forest, containing some deciduous taxa. A number of Amazonian sites record warm temperate rain forest, these assignments responding to the presence of Andean floristic elements within lowland vegetation. There are a number of sites that record tropical dry forest, this being relatively widespread, e.g. on Easter Island, lowland Colombia and the Brazilian interior. Warm temperate rain forest describes the majority of the sites in the Panamanian and southern Costa Rican isthmus with warm mixed forest, being commonly at higher altitudes. This is an area where the comparison between the observed and predicted biomes shows a discrepancy; the possible reasons behind this will be discussed fully. Warm mixed forest is correctly assigned to the highlands of central and southern México as is tropical dry forest on the Yucatán peninsula of southern México. Investigating the correspondence between the pollen-based reconstruction and the potential vegetation for individual biomes provides a check on the methodology, particularly the construction of the matrices. The cool grass/shrubland biome is accurately reconstructed at the majority of sites. The sites that do not match the potential vegetation commonly result from the inclusion of high altitude arboreal pollen, this resulting in assignments of cool mixed forest and cool temperate rain forest. The other common assignment is towards steppe; the dominance of the pollen spectra by Poaceae, and lack of shrubby taxa, result in the assignment to the steppe biome. Indeed, the affinity scores to the cool grass/shrubland and steppe biome at most sites, where one of these biomes is dominant, is normally quite close. For the cool mixed forest biome 66% of sites accurately reconstruct the potential vegetation. The 34% of wrong assignments mainly result in either a reconstruction of cool grass/shrubland, thought to 387 represent possible forest clearance, and the dominance of the vegetation by grassland, or cool temperate rain forest biome due to the numerous shared taxa between these two biomes. 7% of cool temperate rain forest biome reconstructions match the potential vegetation at the site. The remaining % of the sites mainly show either warm mixed forest or warm temperate rain forest assignments. For the tropical dry forest biome some 90% of the sites accurately reflect the potential vegetation at the site. For the remaining % of wrong assignments the common result is towards a cool temperate rain forest or steppe. For the tropical rain forest biome 8% of the sites accurately reflect the potential vegetation. For the sites that do not match, a common reconstruction is warm temperate rain forest. This can be explained by the number of Andean elements being present within lowland tropical forests with a couple of sites reconstructing the closely related biome of tropical seasonal forest. This facet of the pollen data is also exemplified by a number (3%) of the tropical seasonal forest sites recording the warm temperate rain forest biome. Warm evergreen forest is correctly assigned at 80% of the sites. Warm temperate rain forest is assigned correctly at 78% of sites. 7% of the sites that do not reconstruct this biome correctly lead to assignments of tropical rainforest and tropical seasonal forest at low altitudes (<00 m) and cool mixed forest at high altitudes (>00 m). The generally correct biome assignments, in relation to a map of potential vegetation confirm the robustness of our application of the biomisation method to Latin America. Where the match between pollen and potential vegetation reconstructions is relatively low (tropical seasonal forest and warm temperate rain forest), then a common forcing factor, that of high altitude plants presently growing at low altitudes appears important. For other sites where the reconstructed biome does not match the potential vegetation map a series of different explanations, particularly local site-specific factors such as human impact, can be invoked, these will be discussed fully. Taking our modern pollen to potential vegetation calibration, and the design of the matrices that drive it, we reconstruct vegetation at past time intervals with cautious confidence. 388

12 C yr BP biome reconstruction The biomes reconstructed at 6000±00 14 C yr BP (Fig. 8) show relatively small patterns of change compared to the present. 8% of the sites retain the same biome assignment as present (Table 6). In southeastern Brazil, the majority of the sites that were previously assigned to the warm temperate forest biome remain unchanged. A number of sites (e.g. Serra Campos Gerais, Rio São Francisco, Aguads Emendadas) record tropical dry forest at 6000±00 14 C yr BP replacing tropical seasonal forest (Laguna Angel, Laguna Chaplin) record at the present. Sites assigned to tropical rain forest and tropical seasonal forest today mainly remain unchanged at 6000±00 14 C yr BP. Steppe continues to be reconstructed in southeastern Argentina, as today. However, a number of sites had substantially more arboreal components at 6000±00 14 C yr BP than today, for example Empalme Querandíes and Lake Valencia show a transition from steppe to tropical dry forest. An expansion of steppe is recorded at sites previously assigned to cold mixed forest on the Cordillera de los Andes. Unlike sites in southern-most South America that similarly record steppe, these sites also contain significant amounts of Alnus and Podocarpus indicative of parkland at this time. On closer inspection of the affinity scores, sites record an increased affinity to cool temperate rain forest, primarily due to increased amounts of Nothofagus pollen, although this was not sufficiently numerous to produce a cool temperate rain forest assignment. Southernmost South America continues to have a mixture of cool mixed forest, cool grass/shrubland, steppe and cool temperate rain forest biomes, the latter being dominant. Along the southern Andean spine, the assignments do not differ greatly from the modern assignment. There are broadly similar assignments to the present at Colombian sites although there is a slight increase in the number of cool mixed forest and cool temperate rain forest biome assignments relative to cool grass/shrubland of the present day. The sites where this occurs (e.g. La Primevera, and Páramo de Peña Negra) are located at high altitude sites and may reflect either a lowering of the forest line or increased distribution of Andean forest that predates early human impact. Sites 389 in coastal northern South America show a transition to tropical dry forest and tropical seasonal forest from steppe and tropical dry forest respectively, both indicative of a relatively mesic environment. A number of sites on the Yucatán peninsula show a clear distribution of warm evergreen forest at 6000±00 14 C yr BP changing from the warm mixed forest and tropical dry forest reconstruction for the present day. These transitions are not recorded everywhere, for example sites located in the Mexican highlands retain the same biome assignment at the present warm mixed forest C yr BP biome reconstruction Vegetation at ±00 14 C yr BP was substantially different from the present-day, or that reconstructed at 6000±00 14 C yr BP (Fig. 9). The intensity of this vegetation transformation is demonstrated by 82% of the sites change the biome assignment relative to the two previous periods. In Amazonia, tropical seasonal forest and tropical dry forest is recorded instead of tropical rain forest or tropical seasonal forest reconstructed for the present. A site on the present southern Amazonian boundary (Laguna Chaplin) records tropical seasonal forest. Sites in southern South America nearly all sites show a transition from cool mixed forest biomes and cool mixed forest to cool grass/shrubland and cool grassland. However, within this homogenous reconstruction a number of sites have a relatively high affinity to the cool temperate rain forest biome, due to a mix of Donartia and Nothofagus pollen: this explains why the northernmost site in this cluster records cool temperate rain forest. Sites in southeastern Brazil record a transition from tropical dry forest to tropical seasonal forest and cool grass/shrubland. Sites in Amazonia record mainly tropical seasonal forest, warm temperate rain forest or steppe; this combination indicating relatively mesic forest. In the Colombian lowlands, tropical dry forest continues to be assigned whereas Colombian highland locations reflect a marked change from cool temperate rain forest and cool mixed forest to the cool grass/shrubland biome. Sites in Central America show a change from tropical seasonal forest to tropical dry forest, e.g. El Valle. The Mexican highland sites remain unchanged, continuing to support warm mixed forest with the pollen records being 390

13 dominated by Pinus and Quercus. 4 Discussion and conclusions Previous applications of the biomisation method in Africa (Jolly et al., 1998a; Elenga et al., 00), China (Yu et al., 1998, 01), Australia (Pickett et al., 04), Eastern North America (Williams et al., 00), Eurasia (Tarasov et al., 1998a), Europe (Prentice et al., 1996a, b; Tarasov et al., 1998a, b; Elenga et al., 00), Japan (Takahara et al., 01) and Western North America (Thompson and Anderson, 00) demonstrate that technique is able to translate multi-site pollen data to coarse resolution vegetation reconstructions that works well over a range of vegetation types. The Latin American results presented here provide a further test of this ability. The ability of the biomisation method to reconstruct biomes derives in part from the relatively coarse vegetation classification (Fig. 2); which conceals significant intra-biome variation; for example, we do not distinguish subtypes of the warm evergreen forest biome which contains Araucaria in southern and southeastern Brazil and Podocarpus in the northern Andes. The success of the biomisation technique is in part be due to reconstructions being carried out at a regional scale, allowing the methodology to be adapted to the local flora, bioclimatic gradients and pollen spectra. For example, the treatment of Quercus pollen in Latin America is quite different from that in a European context. Similarly, in Africa Podocarpus is assigned to the warm temperate broad-leaved evergreen PFT (Jolly et al., 01), although this taxon is a coniferous needle-leafed tree, in Latin America it is assigned to cool and intermediate temperate conifers. This regional focus also allows the pollen to plant functional type allocations to be based on good ecological information concerned with environmental tolerances to growth limits and an understanding of how representative the pollen is of the surrounding vegetation. This is particularly important as the pollen taxa identified to the generic level (the taxonomic level usually identified to) exhibit considerable plasticity in their growth form and environmental tolerance. For example, within the genus Cordia, commonly a woody shrub of open 391 thorn woodland, of the northern Andes (Cleef and Hooghiemstra, 1984) two species of Cordia are herbs in cerrado (Pereira et al., 1990; Sarmiento, 197), the genus is also present (C. lomatoloba and C. sagotii) in Amazonian terra firme forest and Guyanese lowland rain forest (Steege, 1998). Furthermore the specific nature of pollen production, dispersal and incorporation into a sedimentary environment exhibits considerable variability that is part dependent on site characteristic. All these factors have a bearing on the results and need to be considered in the designing of the input matrices into the biomisation process and interpretation of results. Biomes are mainly accurately reconstructed for the present-day even though large areas of Latin America are covered with vegetation that has been altered by a long history of human land use (Behling, 1996; Binford et al., 1987; Fjeldså, 1992; Gnécco and Mohammed, 1994; Gnécco and Mora, 1997; Marchant et al., 04; Northrop and Horn, 1996). One possible reason for this may relate to the nature of the modern samples. Within our analysis the modern samples are largely derived from sedimentary columns rather than surface trap pollen data, and hence they may stem from the last 00 years and be reflective of a period prior to intensive human-induced change. However, the signal of vegetation clearance does impact on the modern reconstruction as shown by the large number of sites recording cool grass/shrubland, particularly at lower and mid-altitudes that should support cool mixed forest or cool temperate rain forest. These assignments are thought to result from human impact with the pollen spectra being dominated by Poaceae and hence recording more open vegetation. To quantify the nature of this impact, it is possible to tailor the biomisation methodology to include elements of the pollen spectra, such as agricultural and ruderal taxa, that may indicate human impact (Marchant et al., 02). The ability to reconstruct potential, rather than actual vegetation, may also relate to the type of impact; although spatially relatively widespread, forest clearance is often only partial with many localised patches of forest and secondary vegetation remaining. This results in the floristic composition of the remaining vegetation, in palynological terms at least, closely reflecting the original vegetation composition. For example, the forest surrounding the Fúquene-II site is 392

14 a successional type of forest whereas the natural vegetation would be a Andean forest type dominated by Quercus and Weimannia mixed with Croton, Oreopanax and Phyllanthus (Van Geel and Van der Hammen, 1973). In addition to the relatively coarse potential vegetation and biome classification, mapping the highest biome affinity score to each site as a single dot also allows the method to be relativity robust. Although this is suitable for the relatively coarse reconstructions necessitated by the continental/subcontinental scale, when investigating a small area, more information can be preserved from the analysis. Indeed at a regional scale information on sub-dominate biomes can be kept (Marchant et al., 01a), new more defined biomes (Bigelow et al., 03) or at a site-specific scale where the affinity scores in all the biomes can be retained (Marchant et al., 01b, 02b). Despite the overall agreement between potential and reconstructed biomes a number of locations show anomalies. Due to the floristic and structural similarities between warm and cool grasslands (Tarasov et al., 1998a), grass-dominated biomes can be particularly difficult to distinguish from one another. Differentiation is possible by the other plants within steppe and cool grass/shrubland, although there remains a high affinity score to the cool grass/shrubland at low altitudes with the reverse for the steppe biome at high altitudes. Another facet is that some lowland sites show reconstructions of highland biomes, e.g. sites in central Panama and Amazonia recording warm temperate rain forest. This result is driven by the presence of genera that are typical of montane vegetation, e.g. Hedyosmum, Podocarpus and Quercus. A possible explanation for the presence of these highland elements is that they are relictual; relatively isolated today they were previously much more widespread under the glacial climate norm of the Quaternary. This suggestion is supported by the similarity, at a generic level, of the flora in highland Brazil and the northeastern Andes, and the isolated patches of savanna within Amazonian forest and the Brazilian cerrado. Furthermore, it is interesting that the presence of highland elements appears to be greater when the moisture levels are high. For example, within the Chocó Pacific region, where rainfall exceeds 000 mm yr 1, montane elements appear more common than within Amazonia (Gen- 393 try, 1986). Notwithstanding some of the anomalies mentioned, the biomisation method applied to Latin American pollen data can reconstruct large-scale vegetation patterns despite many pollen taxa having different ecological interpretations under different environmental settings (Grabandt, 1980), representation of parent vegetation by pollen likely to be subject to inter-annual variability (Behling et al., 1997b), and tropical vegetation being difficult to reconstruct through pollen assemblages (Bush, 1991; Mancini, 1993; Bush and Rivera, 1998; Behling et al., 1997). These factors demonstrate the importance of basing the input matrices for the biomisation process on all the available ecological information that allowing for the multiple assignment of the pollen taxa to the PFTs. 4.1 Late Quaternary biome changes and palaeoenvironmental interpretations ±00 14 C yr BP Compared to the present, the sites at 6000±00 14 C yr BP record either the same biome or one indicating more xeric vegetation. Dry environmental conditions in southern Brazil extend from the early Holocene until approximately C yr BP when there was an increase in arboreal taxa (Alexandre et al., 1999). Maximum aridity in southeast Brazil was reached between approximately 6000 and C yr BP, prior to the transition to a modern climate (Behling, 1997a). The driest phase in central Brazil is at approximately C yr BP; relatively moist climate conditions similar to today setting in after C yr BP (Ledru, 1993; Marchant and Hooghiemstra, 04). Although fire has been proposed as being responsible for late Holocene variation in the forest/savanna boundary in Brazil (Vernet et al., 1994; Desjardins et al., 1996; Horn, 1993), this relative aridity is thought to reflect an extended dry season during this period (Behling, 1997b). An extended dry season may explain why Araucaria-dominated forest were still restricted in their distribution relative to the modern day, not significantly increasing in range until approximately C yr BP (Behling, 1997a). From our analysis the temporal perspective is missing, hence, we are unable to indicate if 394

15 the vegetation reflects a stable dry period, or a period where there are alternating periods of dry and humid climates linked for example to El Niño activity (Martin et al., 1993; Sifeddine et al., 01). A relatively dry phase is also recorded in northwestern Argentina between 700 and C yr BP (Schäbitz, 1991). Although pollen assemblages do not lend themselves well to distinguishing moisture from temperature changes, stable hydrogen isotope analysis on mosses show the vegetation of southern South America is highly sensitive to changes in moisture regime (Pendall et al., 01). The predominance of steppe in southeastern Argentina agrees with the reconstruction by Prieto (1996): steppe characterising the area between 7000 and C yr BP. Locally high moisture levels at sites closer to the Atlantic Ocean (Prieto, 1996) may explain why sites under strongest maritime influence (Aguads Emendadas, Cerro La China) changes from steppe to tropical dry forest as the local environment is able to support more arboreal taxa. In soutwestern Patagonia a sustained increase in Nothofagus pollen has been detected from around 6800 yr BP thought to result from locally increased moisture levels (Villa-Martinez and Moreno, 07) Locally increased moisture levels in this part of Latin America during the mid Holocene are though to stem from intensification of the southern Westerlies (Gilli et al., 0). Farther west, cool temperate rain forest assignments indicate a similar climatic regime and the maintenance of Valdivian rain forest (Villigran, 1988). A dry phase is also recorded at many Andean sites, for example, in northern Chile desiccation of the Puna ecosystem is recorded between 8000 and C yr BP (Baied and Wheeler, 1993; Villigran, 1988). In lowland Chile, the period of maximum aridity occurred between 9400 and C yr BP with drier than present conditions continuing until C yr BP (Heusser, 1982), this could explain the increased presence of steppe at sites along the southern Andes. On the central Peruvian Andes, a dry warm climate was encountered between 7000 and C yr BP (Hansen, Seltzer and Wright, 1994). δ 18 O measurements from an ice core record taken from highland Peru show that mid-holocene climatic warming and drying was recorded from 80 to 0 14 C yr BP with maximum aridity between 600 to 0 14 C yr BP (Thompson et 39 al., 199). Farther north on the Bolivian Andes, a dry phase is recorded from approximately C yr BP (Abbot et al., 1997). The slight increase in the number of arboreal biome assignments at northern Andean sites can be interpreted as an up-slope shift of forest line. This conforms to the suggestion based on pollen data by van Geel and van der Hammen (1973) that the vegetation zones in the northern Andes were several hundred of meters higher than the present at approximately C yr BP. Relatively dry conditions have also been indicated for lowland Colombia for the mid- Holocene although the onset of dry conditions varied considerably between sites occurring between 600 and C yr BP (Behling et al., 1999). Added complexity is caused by steep environmental gradients associated with non-climatic factors. For example, the presence of the tropical dry forest biome in lowland Colombia, e.g. the catchment of El Piñal, results from a combination of strongly seasonal conditions at present and locally strong edaphic influence (Behling and Hooghiemstra, 1999). Farther north, the assignment of Lake Valencia to the tropical dry forest is in agreement with the site-specific interpretation that more arboreal taxa (Bursera, Piper and Trema) were present after approximately C yr BP due to the onset of a more humid climate (Bradbury et al., 1981): these tropical raingreen taxa indicative of a seasonal climate with relatively dry conditions. This appears to be a regional signal as early Holocene evergreen forests of northern Venezuela were replaced by semi-deciduous elements during the mid-holocene (Leyden, 1984). Enhanced precipitation over Central America being accompanied by a northward shift of the ITCZ, enhanced southerlies and cooler equatorial sea surface temperatures (Harrison et al., 03). Low lake levels in central Panama also indicate that environmental conditions at this period were more xeric (Piperno et al., 1991b; Bush et al., 1992) whereas sites on the Yucatán peninsula show a shift to warm evergreen forest where the warmer conditions that characterise the early Holocene persisted until approximately C yr BP (Brown, 198). This result may stem from locally high moisture levels as a result of maritime influence, a similar mechanism having being proposed to explain a comparable shift in coastal Brazil and Argentina. Despite the majority of the evidence for a mid-holocene dry pe- 396

16 riod, there still remains a debate about the intensity, and even the occurrence, of this. Salgado-Labouriau et al. (1998) suggests that most savanna areas were characterised by increased rainfall between 7000 and C yr BP although there is considerable variation in the timing of the onset of more humid conditions so it may be that such a mesic period falls outside our temporal window. One of the main mechanisms used to explain moisture shifts is fluctuations in the Southern Oscillation and the migration of the ITCZ (Martin et al., 1997). Martin et al. (1997) suggests that during the mid-holocene, the ITCZ was located farther north than its present-day position (Fig. 1) this would produce a summer rainfall deficit and increased winter precipitation; in short greater seasonality. Rather than changes in the median position of the ITCZ, changes in the character of the ITCZ oscillation, such as greater latitudinal range for annual migration, can be invoked to explain vegetation changes (Behling and Hooghiemstra, 01). However, due to the topographical influence of the Andes and the convergence of westerly and easterly winds, the ITCZ has a sinusoidal profile over northern South America (Fig. 1). Therefore, moisture changes over northeastern South America are likely to result from the importance of convective moisture sources; reduced precipitation, particularly in mid latitude western South America, following reduced intensity of westerly climate systems. It is also possible that episodic dry events that presently occur in South America in relation to sea-surface temperature anomalies of the Pacific Ocean (ENSO) were more frequent in the mid-holocene (Markgraf, 1998). This later suggestion may also have led to the increased fire frequency indicated in southeast Brazil (Alexandre et al., 1999). This regression of the forest during the mid-holocene (8000 to C yr BP) in the southern tropical zone of Latin America is opposite to full forest development in Africa (Servant et al., 1993; Jolly et al., 1998a) and this spatial relationship between Latin American and Africa warrants further investigation (Marchant and Hooghiemstra, 04). A particular target for the investigation could be the impact and feedbacks of vegetation changes on climate. For example, large changes in African vegetation about the Sahel are suggested to have been important in influencing Indian monsoon 397 dynamic (Doherty et al., 00). Such a phenomena of vegetation feedbacks on the climate system appears weaker in South America than in Africa although it is likely to have had an impact as yet unqualified. Certainly Latin America would benefit from targeted model applications in the same way that has been applied to Africa (Kubatzki and Claussen, 1998; Doherty et al., 00). This modelling of climate dynamics Latin American represents a special challenge for climate models and modellers (Valdes, 00) primarily due to the steep environmental gradients and rapid transition from one biome to another (Fig. 2) ±00 14 C yr BP The dating of the LGM in Latin America can be problematic (Bush et al., 1990; Hooghiemstra et al., 1992; Ledru et al., 1996, 1998; Sifeddine et al., 01); Late Pleistocene sediments often containing sedimentary gaps at, or about, the LGM (Ledru et al., 1998), that are compounded by slow sedimentation rates. These sedimentary constraints make characterisation of the LGM vegetation highly contentious and have fuelled debates on LGM climates spanning two decades (Hooghiemstra and van der Hammen, 1998; Colinvaux et al., 00; Thomas, 00). Indeed, it has been suggested that some of the sites used in our analysis do not contain a sedimentary record of the LGM (Ledru et al., 1998) although due to application of a 00 year-wide time window, we are able to include some of these sites with contentious sediments. The LGM in Latin America, like most of the tropics, was characterised by a cold dry climate (González et al., 08). Ice caps were present on the southern tip of South America which spread onto the plains and the coastal area (Heine, 199). Evidence from glacial moraines also indicates considerable expansion of Andean glaciers (Hollis and Schilling, 1981; Villagran, 1988; Birkland et al., 1989; Seltzer, 1990; Thouret et al., 1997). Most of southern South America was characterised by an erosional environment; locations that would later accumulate sediments were glaciated, or subject to fluvial activity (Heine, 00). This situation is recorded by ice cores from the high Andes that contain large amounts of dust about the LGM, this being derived from 398

17 surrounding deflating desert areas (Thompson et al., 199). This cold, arid environment is clearly reproduced by the vegetation which shows a transformation from the cool temperate rain forest to the cool grass/shrubland biome. Although Nothofagusdominated forest is thought to have been extirpolated from coastal Chile at the LGM (Hollis and Schilling, 1981), fossil beetle assemblages in basal peat from Puerto Eden (49 S, 74 W) indicate that Nothofagus-dominated forest survived glaciation within the Chilean channels (Ashworth et al., 1991). An earlier date of deglaciation of the Taitao Peninsula indicates migration from Chiloé Island may explain the rapid re-growth of Nothofagus-dominated forest (Lumley and Switsur, 1993). Along the Chilean Pacific coast the present cool evergreen forest was shifted approximately northwards relative to the present day; not as a discrete forest type but as a parkland type vegetation mosaic (Villagran, 1988), not forming closed forests until the early Holocene (Schäbitz, 1994; Heusser, 199). This vegetation is evidenced with the analysis presented here by the northernmost site (Laguna Six Minutes) recording cool temperate rain forest. However, it is unlikely this represents closed forest persisting in the area, trees being present within a woodland/steppe vegetation mosaic (Villagran, 1988). The rate of spreading of this forest into the Holocene would probably have been strongly dependent on the density of the parent plants from the initial seeding fraction (Huntingford et al., 00). The maintenance of cool temperate rain forest taxa, albeit at relatively low levels, may result from high moisture levels as recorded by high lake stands at this time (Markgraf et al., 00). These may reflect outbreaks of polar air and subsequent generation of low-pressure systems in the western Atlantic; combined with lower temperatures this situation would lead to a positive water balance. Indeed, the presence of relatively local moisture sources would have been important at the LGM and allow us to explain regional patterns of biome change outside the influence of the ITCZ migrations (Markgraf et al., 00). Considering the sites along the northern Andes, it is clear from the vegetation that climate was colder during the LGM, reductions up to 12 C may have been reached at very high altitudes (Thompson et al., 199). A substantial temperature depres- 399 sion during the last glacial period is mirrored by a significant impact on the vegetation composition and distribution. From our analysis it is apparent that the tree line was significantly lower at the LGM, concordant with a suggested lowering of vegetation zones by approximately 00 to 00 m relative to the present-day position (Monslave, 198; Wille et al., 01). At lower elevations in western Colombia, a more conservative depression of the vegetation has been suggested from Timbio (Wille et al., 00). Indeed, the spatial character of the cooling and drying in the Neotropics is still under debate (Markgraf, 1993; Colinvaux, 1996; Hooghiemstra and Van der Hammen, 1998; Farrera et al. 1999; Boom et al., 02). Greater temperature change at high altitudes compared with those at low altitudes and at the sea surface (CLIMAP, 1976) can be explained in terms of changes in lapse rate (Bush et al., 1990; Peyron et al., 00; Wille et al., 01) or compression of vegetation belts (Van der Hammen and Absy, 1994). The lapse-rate gradient is partly influenced by atmospheric moisture levels (Barry and Chorley, 1990). As precipitation was reduced at the LGM, an overall steeper lapse rate, particularly at higher altitudes where moisture reductions would have been highest, seems likely (Wille et al., 01). The extent to which lapse-rate changes can be used to explain spatially different signals from the data must be used with caution, particularly as most palaeoclimatic reconstructions have been carried out with some kind of modern analogue-driven transfer function (Farrera et al., 1999). These reconstructions commonly do not take into account non-climatic parameters which would impact on vegetation composition and distribution such as volcanic activity (Kuhry, 1988), fire (Cavelier et al., 1998; Rull, 1999), UV-B insolation (Flenley, 1998) or atmospheric composition, in particular changing CO 2 levels (Woodward and Bond, 04). For example, concentrations of CO 2 reduced to glacial levels (0 ppmv) have been shown to have a very significant impact on tropical vegetation (Jolly and Haxeltine, 1997; Boom et al., 02; Marchant et al., 02b). In south-east Brazil vegetation at the LGM was characterised by tropical dry forest and tropical seasonal forest; this latter vegetation type may have been restricted within deep valleys and along waterways; site-specific records from southeast Brazil indicate 400

18 open grasslands (campo limpo) with forest elements being retained as gallery forest (Behling, 1997a). Some model reconstructions of global vegetation patterns have indicated that there was an increase in warm evergreen forest in Brazil at the LGM at the expense of tropical seasonal forest (Prentice et al., 1993). This pattern of change is supported by the data presented here where plants generally found at high altitudes today were more common in Amazonia at the LGM. Clapperton (1993) used geomorphic data to infer a very sparsely vegetated landscape on the Brazilian Highlands, possibly relating to our reconstruction of steppe for a site in eastern Brazil. However, the cool grass/shrubland biome appears to be a common type of vegetation at this time. Cold climates in eastern South America could have resulted from the incursion of polar cold fronts that would occasionally reach northwards of the equator (Ledru, 1993; Behling and Hooghiemstra, 01). This phenomenon could combine with equatorward shift of polar high-pressure areas and mid-latitude cyclones resulting in displacement and compression of the subtropical anticyclone between mid latitudes westerlies (Dawson, 1992). This climatic regime would result in more restricted migration of the ITCZ and pronounced aridity that would have been compounded by lower sea surface temperatures and associated reduction in atmospheric moisture. At altitudes of approximately 3000 m in northern Peru vegetation about the LGM comprised a mixture of wet and moist montane forest elements with open woodland (Hansen and Rodbell, 199); this vegetation association having no modern analogue. Although the Andes remained relatively moist at the LGM, particularly in the northern part where the concave shape of the mountain chain entrap moisture from the rising air (Fjeldså et al., 1997), it is not certain what occurred in the lowland areas to the east of the Andes (Colinvaux, 1989; Colinvaux et al., 1997; Bush et al., 1990; Thouret et al., 1997). In the Colombian lowlands two sites are characterised by the tropical dry forest biome, this agrees with the suggestion from a pollen study at Rondonia that very open savanna characterised the catchment at the LGM (van der Hammen and Absy, 1994). Similarly, sparse vegetation cover would have been present on the Plateau of Mato Grosso (Servant et al., 1993) and is likely to have extended along the coastal 401 areas of Guyana and Surinam (Wijmstra, 1971) this scenario is supported by our analysis, i.e. a site in lowland Panama recording tropical seasonal forest. Although the majority of the area presently covered by drier types of tropical forest would probably have been replaced by more open woodland at the LGM, environmental changes in savannas at the LGM appear to have been spatially complex. Whether the drier, cooler, conditions resulted in restricted range forest refugia cannot be answered from the available evidence although the vegetation appears heterogeneous as a mosaic of Andean, savanna and tropical rain forest taxa combined. Indeed, this reiterates the suggestion by Colinvaux et al. (01), now widely accepted within the palaeoecological community, that plants responded to Quaternary climate changes as individuals not as biomes. Therefore, to fully investigate vegetation response to climate change is necessary to retain information contained within the affinity scores to the sub-dominant biomes (Marchant et al., 02), or to carry out the analysis at the PFT level. Indeed this approach would allow investigations into which elements of the vegetation were particularly sensitive to environmental change. Expansion of savanna could have been aided by reduced CO 2 concentrations and the resultant competitive advantage attained by C 4 grasses over C 3 plants (Haberle and Maslin, 1999; Marchant et al., 02). Within highland México, warm mixed forest continues to be reconstructed due to the presence of Pinus and Quercus-dominated forests. Although the same biome is recorded at all these periods, it unlikely to be analogous to present day mixed forest; this was characterised by sparsely forested temperate scrub (Binford et al., 1987). Indeed, a strong aridity signal is directly recorded by low lake levels in central México due to reduced northern excursion of the ITCZ, trade wind circulation, and ensuing reduced oceanic-land moisture transfer (Markgraf et al., 00) that would have been reflected in ecosystem response. For example, forest on the Pacific side of the Central America contained a mosaic of high and low altitude forest species; a similarly novel type of forest has also been shown for Mera, Ecuador (Liu and Colinvaux, 198) and Peten, Guatemala (Leyden, 1984). Of the two sites that record the warm evergreen forest biome at this period a site in Guatemala was dominated by Chenopodiaceae, 402

19 Juniperus, Pinus and Quercus. We have presented vegetation reconstructions throughout Latin America at 6000 ±00 14 C yr BP and ±00 14 C yr BP using an objective method based on biomes, constituent PFTs that are described by a set of unique pollen spectra. As a unified methodology has been applied to the pollen data, this reconstruction of biomes provides an objective basis for interpreting large-scale vegetation dynamics, and the environmental controls on these over the Late Quaternary and can be used as a dataset for model-data comparisons at 6000 and yr BP. Changes at 6000±00 14 C yr BP, although relatively small, indicate a transition to more xeric vegetation. The changes at ±00 14 C yr BP are more homogenous and indicative of a cooler, drier climate. These reconstructions are consistent with numerous sitespecific interpretations of the pollen data. The success of the reconstruction has in part been determined by the coarse resolution of biome definitions, and using the most dominant biome for description and interpretation of the results. To develop understanding of vegetation response to environmental change, and possible feedbacks, information that is presently redundant should be retained and the results combined with climate/vegetation modelling initiatives. It is apparent from the relatively sparse coverage, in comparison to Europe and North America, that the Late Quaternary vegetation history of the Neotropical phytogeographical realm remains still relatively poorly resolved despite its importance in model testing, developing biogeographical theory (Tuomisto and Ruokolainen, 1997), and understanding issues concerned with biodiversity and human-environment interactions. It has been shown that environmental change is rarely spatially uniform and as such necessitates an even greater number of sites to determine more precisely this complexity and the driving mechanisms behind this. New sites, located in key areas, combined with the application of a range of proxies of environmental change, are required to refine our understanding of Neotropical ecosystem responses to Late Quaternary climatic variations. 403 Acknowledgements. This research has been funded by the Netherlands Organisation for Scientific Research (NWO) under award 70:198:08 to Henry Hooghiemstra/Rob Marchant. Rob Marchant s recent work to this paper was supported by EU Grant No: EU-MEXT-CT and contributes to the Global Land Project ( This paper contributes to the BIOME 6000-research initiative which is a community-wide collaboration that started in 1994 under four elements of the International Geosphere-Biosphere Program (IGBP-GAIM, IGBP-DIS, IGBP-GCTE and IGBP-PAGES) with the aim to create fully documented pollen and plant macrofossil data sets for 6000 and C yr BP. Dominic Jolly was instrumental in developing and applying these techniques in Africa and subsequent application to different geographical regions indeed the Latin American application presented here would not have been possible without this massive contribution an encouragement throughout the development of this work. In particular we thank Sandra Diaz, Liz Pickett, Colin Prentice and Bob Thompson for comments on earlier drafts of this manuscript. Ary Teixeira de Oliveira-Filho and Olando Rangel are particularly thanked for comments on the ecology of the Latin American taxa. Kirsten Sickel, Gerhard Boenish and Silvana Schott are thanked for help in producing the figures and archiving the results. Particular thanks must go to Eric Grimm, John Keltner and Vera Markgraf for their energies in establishing, and developing, the Latin American Pollen Database. At the very centre of this work are all those palynologists who have supplied data to the LAPD, or specifically to be used for this paper; without their efforts such syntheses would not be possible. The publication of this article is financed by CNRS-INSU. References Abbot, M. B., Sletzer, G., Kelts, K. R., and Southon, J.: Holocene paleohydrology of the tropical Andes from lake records, Quaternary Res., 47, 70 80, Absy, M. L.: A palynolgical study of Holocene sediments in the Amazon basin, Ph.D. thesis, University of Amsterdam, The Netherlands,

20 30 Absy, M. L., Cleef, A., Fournier, M., Servant, M., Siffedine, A., Da Silva, M. F., Soubies, F. Suguio, K., Turcq, B., and van der Hammen, T.: Mise en évidence de quatre phases d ouverture de la forêt dense dans la sud-east de l Amazonie au cours des dernières années, Première comparison avec d autres regions tropicales, Competes Rendues Academie Science Paris, 313, , Alexandre, A., Meunier, J. D., Mariotti, A., and Soubies, F.: Late Holocene phytolith and carbonisotope record from a latosol at Salitre, South-Central Brazil, Quaternary Res., 1, , Almeida, L.: Vegetación, fitogeographica y paleoecologíca del zacatonal alpine y bosques montanos de la región central de México, Ph.D. thesis, University of Amsterdam, Armesto, J. J., Villágran, C. Aravena, J. C., Pérez, C., Smith-Ramirez, C., Cortés, M., and Hedin, L.: Conifer forests of the Chilean coastal ranges, Ecology of Southern Conifers, edited by: Enright, N. J. and Hill, R. S., Melbourne University Press, , 199. Ashworth, A. C. and Markgraf, V.: Climate of the Chilean Channels between and 000 yr BP based on fossil pollen and beetle analysis, Rev. Chil. Hist. Nat., 62, 61 74, Ashworth, A. C., Markgraf, V., and Villagran, C.: Late Quaternary climatic history of the Chilean channels based on fossil pollen and beetle analyses, with an analysis of the modern vegetation and pollen rain, J. Quaternary Sci., 6, , Argollo, J. and Mourguiart, P.: Late Quaternary climate history of the Bolivian Altiplano, Quatern. Int., 72, 37 1, 00. Baied, C. A. and Wheeler, J. C.: Evolution of high Andean Puna ecosystems: environment, climate, and cultural change over the last years in the central Andes, Mt. Res. Dev., 13, 14 6, Barry, P. and Chorley, L.: Mountain weather and climate, Methuen, London and New York, 364 pp., Beard, J. S.: The classification of tropical American vegetation, J. Ecol., 36, 89 0, 19. Behling, H.: Untersuchungen zur spätpleistozänen und Holozänen Vegetations und Klimageschichte der tropische Kustenwälder und der Araukarienwälder in Santa Catarina (Süd Brazilien), Dissertations Botanicae Band 6, J. Cramer, Berlin Stuttgart, 149 pp., Behling, H.: First report on new evidence for the occurrence of Podocarpus and possible human presence at the mouth of the Amazon during the Late-glacial, Veg. Hist. Archaeobot.,, , Behling, H.: Late Quaternary vegetation, climate and fire history from the tropical mountain region of Morro de Itapeva, SE Brazil, Palaeogeogr. Palaeocl., 129, , 1997a. Behling, H.: Late Quaternary vegetation, climate and fire history of the Araucaria forest and campos region from Serra Campos Gerais, Paraná State (South Brazil), Rev. Palaeobot. Palyno., 97, 9 121, 1997b. Behling, H., Berrio, J. C., and Hooghiemstra, H.: Late Quaternary pollen records from the middle Caquetá River basin in central Colombian Amazon, Palaeogeogr. Palaeocl., 14, , Behling, H. and Hooghiemstra, H.: Late Quaternary palaeoecology and palaeoclimatology from pollen records of the savannas of the Llanos Orientales in Colombia, Palaeogeogr. Palaeocl., 139, 1 267, Behling, H. and Hooghiemstra, H.: Environmental history of the Colombian savannas of the Llanos Orientales since the Last Glacial Maximum from lake records El Piñal and Carimagua, J. Paleolimnol., 21, , Behling, H., and Hooghiemstra, H.: Holocene Amazon rain forest-savanna dynamics and cli- matic implications: high-resolution pollen record from Laguna Loma Linda in eastern Colombia, J. Quaternary Sci.,, , 00. Behling, H. and Hooghiemstra, H.: Neotropical savanna environments in space and time: Late Quaternary inter-hemispheric comparisons, in: Inter-hemispheric climate linkages, edited by: Markgraf, V., Academic Press, , 01. Behling, H., Hooghiemstra, H., and Negret, A. J.: Holocene history of the Chocó rain forest from Laguna Piusbi, Southern Pacific Lowlands of Colombia, Quaternary Res., 0, , Behling, H., Keim, G., Irion, G., Junk, W., and Nunes de Mello, J.: Holocene changes in the Central Amazon Basin inferred from Lago Calado (Brazil), Palaeogeogr. Palaeocl., 173, 87 1, 01. Behling, H. and Lima da Costa, M.: Holocene vegetational and enviornmental changes from the Lago Crispim record in northeastern Para State, eastern Amazonia, Rev. Palaeobot. Palyno., 114, 14, 01. Behling, H., Negrelle, R. B., and Colinvaux, P.: Modern pollen rain from the tropical Atlantic rain forest, Reserve Volta Velha, South Brazil, Rev. Palaeobot. Palyno., 97, , Behling, H., Negret, A. J., and Hooghiemstra, H.: Late Quatrnary vegeteational and climatic change in the Popayan region, southern Colombian Andes, J. Quaternary Sci., 13, 43 3, 406

21 Bennet, K. D., Haberle, S., and Lumley, S. H.: The Late Glacial-Holocene transition in southern Chile, Science, 290, 3 327, 00. Berrio, J. C., Hooghiemstra, H., Behling, H., and Van der Borg, K.: Late Holocene history of savanna gallery forest from Carimagua area, Colombia, Rev. Palaeobot. Palyno., 111, , 00. Berrio, J. C., Behling, H., and Hooghiemstra, H.: Tropical rain forest history from the Colombian Pacific area: a 40-yr pollen record from Laguna Jotaordó, Holocene,, , 00. Berrio, J. C., Hooghiemstra, H., Behling, H., Botero, P., and Van der Borg, K.: Late Quaternary savanna history of the Colombian Llanos Orientales from Lagunas Chenevo and Mozambique: a trasect synthesis, Holocene, 12, 3 48, 02. Berrio, J. C., Boom, A., Botero, P., Herrera, L., Hooghiemstra, H., Romero, F., and Sarmiento, G.: Multi-interdisciplinary evidence of The Holocene history of a cultivated floodplain area in the wetlands, Veg. Hist. Archaeobot.,, , 02. Berrio, J. C., Hooghiemstra, H., Marchant, R., and Rangle, O.: Late glacial and Holocene history of dry forest in the Cauca Valley from sites Quilichao and La Teta, J. Quaternary Sci., 17, , 02. Bigelow, N. H., Brubaker, L. B., Edwards, M. E., Harrison, S. P., Prentice, I. C., Anderson, P. M., Andreev, A. A., Bartlein, P. J., Christensen, T. R., Cramer, W., Kaplan, J. O., Lozhkin, A. V., Matveyeva, N. V., Murray, D. V., McGuire, A. D., Razzhivin, V. Y., Ritchie, J. C., Smith, B., Walker, D. A., Gajewski, K., Wolf, V., Holmqvist, B. H., Igarashi, Y., Kremenetskii, K., Paus, A., Pisaric, M. F. J., and Vokova, V. S.: Climate change and Arctic ecosystems I. Vegetation changes north of N between the last glacial maximum, mid-holocene and present, J. Geophys. Res., 8, 1 17, 03. Binford, M. W., Brenner, M., Whitmore, T. J., Higuera-Gundy, A., Deevey, E. S., and Leyden, B. W: Ecosystems, palaeoecology and human disturbance in subtropical and tropical America, Quaternary Sci. Rev., 6, 1 128, Birkland, P. W., Rodbell, D. T., and Short, S. K.: Radiocarbon dates on deglaciation, Cordillera Central, Northern Peruvian Andes, Quaternary Res., 32, , Björck, S., Håkansson, H., Olsson, S., Barnekow, L., and Janssens, J.: Palaeoclimatic studies in South Shetland Islands, Antarctica, based on numerous stratigraphic variables in lake sediments, J. Paleolimnol., 8., , Boom, A., Marchant, R. A., Hooghiemstra, H., and Sinninghe Damste, J. S.: CO 2 and temperature-controlled altitudinal shifts of C 4 - and C 3 -dominated grasslands allow reconstruction of ρco 2, Palaeogeogr. Palaeocl., 177, 29 4, 02. Braconnot, P., Otto-Bliesner, B., Harrison, S., Joussaume, S., Peterchmitt, J.-Y., Abe-Ouchi, A., Crucifix, M., Driesschaert, E., Fichefet, Th., Hewitt, C. D., Kageyama, M., Kitoh, A., Laîné, A., Loutre, M.-F., Marti, O., Merkel, U., Ramstein, G., Valdes, P., Weber, S. L., Yu, Y., and Zhao, Y.: Results of PMIP2 coupled simulations of the Mid-Holocene and Last Glacial Maximum - Part 1: experiments and large-scale features, Clim. Past, 3, , 07, Bradbury, J. P., Leyden, B., Salgado-Labouriau, M. L., Lewis Jr. W. M., Schubert, C., Binford, M. W., Schubert, C. Binford, M. W., Frey, D. G., Whitehead, D. R., and Weibezahn, F. H.: Late Quaternary environmental History of Lake Valencia, Venezuela, Science, 214, , Brown, B.: A summary of Late Quaternary pollen records from México west of the Isthmus of Tehuantepec, Pollen records of Late Quaternary North American Sediments, edited by: Bryant, V. M. and Holloway, R. G., American Association of Stratigraphic Palynologists Foundation, Dallas, Texas, 342 pp., 198. Brown, S. and Lugo, A. E.: Tropical secondary forests, J. Trop. Ecol., 6, 1 32, 199. Bush, M.: Modern pollen-rain data from South and Central America: a test of the feasibility of fine-resolution lowland tropical palynology, Holocene, 1, , Bush, M.: Neotropical plant reproductive strategies and fossil pollen representation, Am. Nat., 14, , 199. Bush, M. B. and Colinvaux, P.: A 7000-year pollen record from the Amazon lowlands, Ecuador, Vegetatio, 76, 141 4, Bush, M., Colinvaux, P., Wiemann, M. C., Piperno, D. R., and Liu, K.: Late Pleistocene tem- perature depression and vegetation change in Ecuadorian Amazonia, Quaternary Res., 34, , Bush, M. B., Miller, M., de Oliveira, P. E., and Colinvaux, P.: Two histories of environmental change and human disturbance in eastern lowland Amazonia, Holocene,, 43 3, 00. Bush, M., Piperno, D. O., Colinvaux, P., Oliveria, P. E., Krissek, L. A., Miller, M. C., and Rowe, W. E.: A yr palaeoecological profile of a lowland tropical lake in Panama, Ecol. Monogr., 62, 1 27,

22 30 Bush, M. and Rivera, M.: Pollen dispersal and representation in a neotropical rain forest, Global Ecol. Biogeogr., 7, , Björck, S., Håkansson, S. H., Olsson, S., Barnekow, L., and Janssens, J. A.: Palaeoclimatic studies in South Shetland Islands, Antarctica, based on numerous stratigraphic variables in lake sediments, J. Paleolimnol., 8, , Byrne, R. and Horn, S. P.: Prehistoric agriculture and forest clearance in the Sierra de los Tuxtlas, Veracruz, México, Palynolgy, 13, , Cavelier, J., Aide, T. M., Santos, C., Eusse, A. M., and Dupuy, J. M.: The savannization of moist forests in the Sierra Nevada de Santa Marta, Colombia, J. Biogeogr.,, , Cerverny, R. S.: Present climates of South America, in: Climates of the Southern Continents: Present, Past and Future, edited by: Hobbs, J. E., Lindesay, J. A., and Bridgeman, H. A., J. Wiley and Sons, Chichester, , Clapperton, C. M.: Quaternary geology and geomorphology of South America, Elsevier, Amsterdam, Claussen, M.: On coupling global climate models with climate models, Climate Res., 4, 3 221, Claussen, M. and Esch, M.: Biomes computed from simulated climatologies, Clim. Dynam., 9, , Cleef, A. M. and Hooghiemstra, H.: Present vegetation of the area of the high plain of Bogotá, in: Vegetation and Climatic History of the High Plain of Bogotá, Colombia, edited by: Hooghiemstra, H., Ganter Verlag, Vaduz, Dissertationes Botanicae, 79, 1 368, CLIMAP Project Members: The surface of ice-age Earth, Science, 191, , Colinvaux, P. A.: Ice-age Amazon revisited, Nature, 340, , Colinvaux, P. A.: Quaternary Environmental History and Forest Diversity in the Neotropics, in: Evolution and Environment in Tropical America, edited by: Jackson, B. C., Budd, A. F., and Coates, A. G., The University of Chicago Press, Chicago, London, 482 pp., Colinvaux, P. A., Bush, M., Steinitz-Kannan, M., and Miller, M. C.: Glacial and postglacial pollen records from the Ecuadorian Andes and the Amazon, Quaternary Res., 48, 69 78, Colinvaux, P. A., De Oliveira, P. E., Moreno, J. E., Miller, M. C., and Bush, M.: A long pollen record from Lowland Amazonia: Forest and cooling in glacial times, Science, 274, 8 88, Colinvaux, P. A., de Oliveira, P. E., and Bush, M. B.: Amazonian and neotropical communities on glacial time-scales: the failure of the aridity and refuge hypothesis, Quaternary Sci. Rev., , , 00. Colinvaux, P. A., Frost, M., Frost, I., Liu, K B., and Steinitz-Kannan, M.: Three pollen diagrams of forest disturbance in the western Amazon basin, Rev. Palaeobot. Palyno.,, 83 99, Cuatrecasas, J. and Barreto, A. J.: Pàramos, Villegas editores, Bogotá, Colombia, De Oliveira, P. E.: A palynological record of Late Quaternary vegetation and climatic change in southeastern Brazil, Ph.D. thesis, Ohio State University, Colombus, OH, De Oliveira, P. E., Barreto, A. M. F., and Suguio, K.: Late Pleistocene/Holocene climatic and vegetational history of the Brazilian caatinga: the fossil dunes of the middle Sao Francisco River, Palaeogeogr. Palaeocl., 2, , Dawson, A. G.: Ice Age Earth: Late Quaternary Geology and Climate, Routledge, Desjardins, T., Carneiro-Filho, A., Mariotti, A., Chauvel, A., and Girardin, C.: Changes of the forest-savanna boundary in Brazilian Amazonia during The Holocene revealed by stable isotope ratios of soil organic carbon, Oecologia, 8, , Dominquez, G., Rojo-Vazquez, J. R., Galvan-Pina, V., and Aguilar-Palomino, B.: Changes in the structure of a coastal fish assemblage exploited by a small scale gillnet fishery during an El Niño La Niña event, Estuar. Coast. Shelf S., 1, , 00. Dov Par, F.: Sooretama: The Atlantic rain forest of Brazil, SPB Academic Publishing, The Hague, Doherty, R., Kutzbach, J., Foley, J., and Pollard, D.: Fully coupled climate/dynamical vegetation model simulations over northern Africa during the mid-holocene, Clim. Dynam., 16, 61 73, 00. Duivenvoorden, J. F. and Cleef, A. M.: Amazonian savanna vegetation on the sandstone near Araracuara, Colombia, Phytocoenesis, 24, , Edwards, M. E., Anderson, P. M., Brubaker, L. B., Ager, T., Andreev, A. A., Bigelow, N. H., Cwynar, L. C., Eisner, W. R., Harrison, S. P., Hu, F-S., Jolly, D., Lozhkin, A. V., MacDonald, G. M., Mock, C. J., Ritchie, J. C., Sher, A. V., Spear, R. W., Williams, J., and Yu, G.: Pollenbased biomes for Beringia , 6000 and 0 14 C yr BP, J. Biogeogr., 27, 21 4, 00. Eidt R. C.: The climatology of South America, in: Biogeography and ecology in South America, edited by: Fittkau, E. J., Junk, The Hague, 1, 4 81, Eiten, G.: The cerrado vegetation of Brazil, Bot. Rev., 2, 2 341, Elenga, H., Peyron, O., Bonnefille, R., Prentice, I. C., Jolly, D., Cheddadi, R., Guiot, J., Andrieu, V., Bottema, S., Buchet, G., de Beaulieu, J. L., Hamilton, A. C., Maley, J., Marchant, R., 4

23 30 Perez-Obiol, R., Reille, M., Riollet, G., Scott, L., Straka, H., Taylor, D., Van Campo, E., Vincens, A., Laarif, F., and Jonson, H.: Pollen-based reconstruction for southern Europe and Africa years ago, J. Biogeogr., 27, , 00. Ellis, E. C. and Ramankutty, N.: Putting people in the map: anthropgenic biomes of the world, Front. Ecol. Environ., 6, 18, 08. Farrera, I., Harrison, S. P., Prentice, I. C., Ramstein, G., Guiot, J., Bartlein, P. J., Bonnefille, R., Bush, M., Cramer, W., von Grafenstein, U., Holmgren, K., Hooghiemstra, H., Hope, G., Jolly, D., Lauritzen, S. E., Ono, Y., Pinot, S., Stute, M., and Gu, Y.: Tropical climates at the last glacial maximum: a new synthesis of terrestrial palaeoclimate data. I. Vegetation, lake-levels and geochemistry, Clim. Dynam., 11, , Flenley, J. R.: Tropical forests under the climates of the last years, Climate Change, 39, , Flenley, J. R. and King, S. M.: Late Quaternary pollen records from Easter Island, Nature, 307, 47 0, Flenley, J. R., King, S. M., Jackson, J., Chew, C., Teller, J. T., and Prentice, M. E.: The late Quaternary vegetation and climatic history of Easter Island, J. Quaternary Sci., 6, 8 1, Fjeldså, J. A.: Un analisis biogeografico de la avifauna de los bosques de quenoa (Polylepis) de los Andes y su relevancia para establecer prioridades de conservacion, Memorias del Museo de Historia Natural, U.N.M.S.M. (Lima), 21, 7 221, Fjeldså, J. A.: The avifauna of the Polylepis woodlands of the Andean highlands: the efficiency of basing conservation priorities on patterns of endemism, Bird Conserv. Int., 3, 3, Fjeldså, J. A., Ehrlich, D., Lambin, E., and Prins, E.: Are biodiversity hot spots correlated with current ecoclimatic stability?, A pilot study using the NOAA-AVHRR remote sensing data, Biodivers. Conserv., 6, 399 4, Foley, J. A., Prentice, I. C., Ramankutty, N., Levis, S., Pollard, D., Sitch, S., and Haxeltine, A.: An integrated biosphere model of land surface processes, terrestrial carbon balance and vegetation dynamics, Global Biogeochem. Cy.,, , Gilli, A., Anselmetti, F. S., Ariztegui, D., Beres, M., McKenzie, J. A., and Markgraf, V.: Seismic stratigraphy, buried beach ridges and contourite drifts: the Late Quaternary history of the closed Lago Cardiel basin, Argentina (49 S), Sedimentology, 2, 1 22, 04. Gentry, A. H.: A field guide to the families and genera of woody plants of South America (Colombia, Ecuador, Perú), Conservation International, Washington, D.C., Gentry, A. H.: Species richness and floristic composition of the Choco region plant communities, Caldasia, 40, 71 91, Godinez-Domínquez, M., Rojo-Vazquez, J. R., Galvan-Pina, V., and Aguilar-Palomino, B.: Changes in the structure of a coastal fish assemblage exploited by a small scale gillnet fishery during an El Niño-La Niña event, Estuar. Coast. Shelf S., 1, , 00. Godley, E. J. and Moar, N. T.: Vegetation and pollen analysis of two bogs on Chiloé, New Zeal. J. Bot., 11, 268, Gnécco, C.: An archaeological perspective on the Pleistocene/Holocene boundary in northern South America, Quatern. Int., 3, 3 9, Gnécco, C. and Mohammed, A.: Technologia de cazadores-recolectores subandinos: Analisis functional y organización tecnologica, Review of Colombian Anthropology, 31, 6 31, Gnécco, C. and Mora, S.: Early occupations of the tropical forests of northern South America by hunter-gatherers, Antiquity, 71, , Gentry, A. H.: A field guide to the families and genera of woody plants of South America (Colombia, Ecuador, Peru), Conservation International, Washington, D.C., Grabandt, R. J.: Pollen rain in relation to arboreal vegetation in the Colombian Cordillera Oriental, Rev. Palaeobot. Palyno., 29, 6 147, Graf, K.: Palinologiadel cuaternario reciente en los Andes del Ecuador, del Peru, y de Bolivia, Boletin Servicio Geologico Bolivia, 4, 69 91, Graf, K.: Pollen diagramme aus den Anden: Eine Synthese zur Klimageschichte und Vegeta- tionsentwicklung seit der letzen Eiszeit, University of Zurich, Switzerland, Physiche Geographie, 34, 72 94, Graherr, G.: Farbatlas Ökosysteme der Erde: natürliche, naturnahe bund künstliche Land- Ökosysteme aus geobotanischer Sicht, Ulmer, Stuttgart, Germany, Hansen, B. C. S. and Rodbell, D. T.: A glacial/holocene pollen record from the eastern Andes of northern Peru, Quaternary Res., 44, , 199. Hansen, B. C. S., Seltzer, G. O., and Wright Jr., H. E.: Late Quaternary vegetational change in the central Peruvian Andes, Palaeogeogr. Palaeocl., 9, , Haberle, S. and Maslin, M.: Late Quaternary vegetation and climate change in the Amazon Basin based on a year pollen record from the Amazon fan, ODP site 932, Quaternary Res., 1, 27 38, Harley, R. M.: Introduction to the vegetation, in: Flora of the Pico das Almas, Chapada Diamantina, Bahia, Brazil, edited by: Stannards, B. L., Royal Botanic Gardens, Kew,

24 30 Harrison, S. P., Kutzbach, J. E., Liu, Z., Bartlein, P. J., Otto-Bliesner, B., Muhs, D., Prentice, I. C., and Thompson, R. S.: Mid-Holocene climates of the Americas: a dynamical response to changed seasonality, Clim. Dynam.,, , 03. Haxeltine, A. and Prentice, I. C.: BIOME3: an equilibrium terrestrial biosphere model based on ecophysiological constraints, resource availability, and competition among plant functional types, Global Biogeochem. Cy.,, , Heine, K. K.: Late Quaternary glacial sequences in Bolivia, Ecuador and México data and palaeoclimatic implications, Abstracts, 14th INQUA Congress, Berlin, 199. Heine, K. K.: Tropicl South America during the Last Glacial maximum: evidence from glacial, periglacial and fluvial records, Quatern. Int., 72, 7 21, 00. Helmens, K. F. and Kuhry, P.: Middle and late Quaternary vegetational and climatic history of the Páramo de Agua Blanca (Eastern Cordillera Colombia), Palaeogeogr. Palaeocl., 6, , Heusser, C. J.: Quaternary palynology of Chile, Quaternary of South America and the Antarctic Peninsula, 1, 22, Heusser, C. J.: Three late Quaternary pollen diagrams from Southern Patagonia and their palaeoecological implications, Palaeogeogr. Palaeocl., 118, 1 24, 199. Heusser, C., Denton, G. H., Hauser, A., Anderson, B., and Lowell, T.: Quaternary pollen records from the Archipelago de Chiloe in the context of glaciation and climate, Rev. Geol. Chile, 22, 46, 199. Heusser, C. Lowell, T., Heusser, L., Moreira, A., and Moreira, S. M.: Pollen sequence from the Llanquihue glaciation in marine Oxygen Isotope Stage 4 2, J. Quaternary Sci.,, 1 1, 00. Hollis, J. H. T. and Schilling, D. H.: Late Wisconsin-Weichselian mountain glaciers and small ice caps, in: The last great ice sheets, edited by: Denton, G. H. and Hughes, T. J., Wiley, New York, 484 pp., Hooghiemstra, H.: Vegetation and climate history of the high plain of Bogotá, Colombia: a continuous record of the last 3. million years, Ganter Verlag, Vadiz, Hooghiemstra, H. and Cleef, A. M.: Development of Vegetational and Climatic Sequence of the Area of the High Plain of Bogotá, in: Vegetation and climate history of the high plain of Bogotá, Columbia: a continuous record of the last 3. million years, edited by: Hooghiemstra, H., Ganter Verlag, Vadiz, Hooghiemstra, H., Cleef, A. M., Noldus, G. W., and Kappelle, M.: Upper Quaternary vegetation dynamics and palaeoeclimatology of the La Chonta bog area (Cordillera de Talamanca, Costa Rica), J. Quaternary Sci., 7, 2, Hooghiemstra, H. and van der Hammen, T.: Neogene and Quaternary development of the neotropical rain forest: the forest refugia hypothesis, and a literature overview, Earth-Sci. Rev., 44, 147 3, Horn, S. P.: Prehistoric fires in the highlands of Costa Rica: sedimentary charcoal evidence, Rev. Biol. Trop., 37, , Horn, S. P.: Postglacial vegetation and fire history in the Chirripó Páramo of Costa Rica, Quaternary Res., 40, 7 116, Hück, K.: Die waldgeographische Regionen und Unterregionen von Südamerika (with map 1: ), Geographische Taschenbuch, Huntingford, C., Cox, P. M., and Lenton, T. M.: Contrasting response of a simple terrestrial ecosystem model to global change, Ecol. Model., 134, 41 8, 00. International Plant Names Project: International Plant Names Index, available at: ipni.org(last access: September 08), Islebe, G. and Hooghiemstra, H.: Recent pollen spectra of highland Guatemala, J. Biogeogr., 22, 91 99, 199. Islebe, G. and Hooghiemstra, H.: Vegetation and the Climate History of Montane Costa Rica since the Last Glacial, Quaternary Sci. Rev., 16, , Islebe, G., Hooghiemstra, H., and van der Borg, K.: A cooling event during the Younger Drays Chron in Costa Rica, Palaeogeog., Palaeoclim., Palaeoecol., 117, 73 80, 199a. Islebe, G., Hooghiemstra, H., and van der Borg, K.: The Younger Drays c in the Cordillera de Talamanca, Costa Rica, Geol. Mijnbouw, 74, , 199b. Islebe, G., Hooghiemstra, H., Brenner, M., Curtis, J. H., and Hodell, D. A.: A Holocene vegeta- tion history from lowland Guatemala, Holocene, 6, , Jolly, D. and Haxeltine, A.: Effect of low glacial atmospheric CO 2 on tropical African montane vegetation, Science, 276, , Jolly, D., Prentice, I. C., Bonnefille, R., Ballouche, A., Bengo, M., Brenac, P., Buchet, G., Burney, D., Cazet, J-P., Cheddadi, R., Edorh, T., Elenga, H., Elmoutaki, S., Guiot, J., Laarif, F., Lamb, H., Lezine, A-M., Maley, J., Mbenza, M., Peryon, O., Reille, M., Reynaud-Farrera, I., Riollet, G., Ritchie, J. C., Roche, E., Scott, L., Ssemmanda, I., Straka, H., Umer, M., Van Campo, E., Vilimumbalo, S., Vincens, A., and Waller, M.: Biome reconstruction from pollen and plant 414

25 30 macrofossil data for Africa and the Arabian peninsula at 0 and 6 ka, J. Biogeogr.,, 997 0, 1998a. Jolly, D., Harrison, S. P., Damnati, B., and Bonnefille, R.: Simulated climate and biomes of Africa during the Late Quaternary: comparisons with pollen and lake status data, Quaternary Sci. Rev., 17, , 1998b. Kahn, F. and de Granville, J. J.: Palms in the forest ecosystems of Amazonia, Springer Verlag, 226 pp., Germany. Kappelle, M.: Structural and floristic differences between western Atlantic and moist Pacific montane Myrsine-Quercus forest in Costa Rica, Academic Press, 483 pp., Kappelle, M.: Ecology of mature and recovering Talamancan Montane Quercus forests, Costa Rica, Ph.D. thesis, University of Amsterdam, The Netherlands, 273 pp., 199. Kershaw, A. P. and McGlone, M.: The Quaternary history of the southern conifers, in: Ecology of Southern Conifers, edited by: Enright, N. J. and Hill, R. S., Melbourne University Press, 342 pp., 199. Kewensis: Index Kewensis version 2.0, Developed by systems simulations Ltd., Oxford Univer- sity Press, CD rom, Kubatzki, C. and Claussen, M.: Simulation of the global biogeophysical interactions during the Last Glacial Maximum, Clim. Dynam., 14, , Kuhry, P.: A paleobotanical and palynological study of Holocene peat from the El Bosque Mire, located in a volcanic area of the Cordillera Central of Colombia, Rev. Paleobot. Polyno.,, 19 72, 1988a. Kuhry, P.: Palaeobotantical-palaeoecological studies of tropical high Andean peatbog sections (Cordillera Oriental, Colombia), Dissertationes Botanicae 116, J. Cramer, Berlin-Stuttgart, 241 pp., Kuhry, P., Salomons, J. B., Riezebos, P. A., and Van der Hammen, T.: Paleoecologia de los ultimos anos en el area de la Laguna de Otun-El Bosque, in: Studies on Tropical Andean Ecosystem, edited by: van der Hammen, T., Perez, P. A., and Pinto, P., La Cordillera Central Colombiana transecto Parque Los Nevados, Cramer, Vaduz, 1, , Latin American Pollen Database (LAPD), available at: last access: September 04. Ledru, M.-P.: Modifications de la vegetation du Brazil entre la dernieier epoque glaciaire et línterglaciaire ectuel, Comptes Rendus de l Academie des Sciences Paris, 314, , Ledru, M.-P.: Late Quaternary environmental and climatic changes in Central Brazil, Quaternary Res., 39, 90 98, Ledru, M.-P., Bertaux, J., and Sifeddine, A.: Absence of Last Glacial Maximum records in lowland tropical forests, Quaternary Res., 49, , Ledru, M.-P., Behling, H., Fournier, M. L., Martin, L., and Servant, M.: Localisation de la fôret d Acaucaria du Brazil au cours de l Holocène, Implications paleoclimatiques, C. R. Acadamie Science Paris, 317, 17 21, Ledru, M.-P., Corderio, R. C., Dominquez, J. M. L., Martin, L., Mourguiart, P., Sifeddine, A., and Turcq, B.: Late-glacial cooling in Amazonia inferred from pollen at Lagoa de Caco, northern Brazil, Quaternary Res.,, 47 6, 01. Ledru, M.-P. and Lorscheitter, M. L.: Vegetation dynamics in southern and Central Brazil during the last 000 yr BP, Rev. Palaeobot. Palyno., 99, , Ledru, M.-P., Soares Braga, P. I., Soubies, F., Fournier, M. L., Martin, L., Suguio, K., and Turcq, B.: The last years in the Neotropics (southern Brazil): evolution of vegetation and climate, Palaeogeogr. Palaeocl., 123, 239 7, Leemans, R. and Cramer, W. P.: The IISA database for mean monthly values of temperature, precipitation and cloudiness on a global terrestrial grid, Rep IIASA Research RR-91-18, International Institute for Applied Systems Analysis, Laxenburg, Austria, Leyden, B. W.: Guatemalan forest synthesis after Pleistocene aridity, P. Natl. Acad. Sci. USA, 81, , Leyden, B. W.: Late Quaternary aridity and Holocene moisture fluctuations in the Lake Valencia Basin, Venezuela, Ecol., 66, , 198. Leyden, B. W.: Man and climate in Maya lowlands, Quaternary Res., 28, , Leyden, B. W., Brenner, M., Hodell, D. A., and Curtis, J. H.: Late Pleistocene climate in the central American lowlands, in: Climate Change in Continental Isotopic Records, edited by: Swar, P. K., American Geophysical Union Geophysical Monograph, Washington, DC, USA, 78, 374 pp., Leyden, B. W., Brenner, M., Hodell, D. A., and Curtis, J. H.: Orbital and internal forcing of the climate on the Yucatán peninsula for the past ca 36 ka, Palaeogeogr. Palaeocl., 9, 193 2, Leyden, B. W., Brenner, M., Whitmore, T., Curtis, J. H., Piperno, D. R., and Dahlin, B. H.: A record of long and short-term climatic variation from northwest Yucatán: Cenote San Jose Chulchaca, in: The managed mosaic: Ancient Maya agriculture and resource use, edited by: 416

26 30 Fedick, S. L., University of Utah Press, USA, 424 pp., 199. Liu, K. and Colinvaux, P.: Forest changes in the Amazon Basin at the LGM, Nature, 318, 6 77, 198. Lozano-Garcia, M. S. and Ortega-Guerroero, B.: Palynological and magnetic susceptibility records of Lake Chalco, central México, Palaeogeogr. Palaeocl., 9, , Lozano-Garcia, M. S., Ortega-Guerroero, B., Caberallero-Mirandan, M., and Urrutia- Fucugauchi, F.: Late Holocene Paleoenvironments of Chalco Lake, Central México, Quaternary Res., 40, , Lozano-Garcia, M. S. and Xelhuantzi-Lopéz, M. S.: Some problems in the late Quaternary pollen records of Central México: Basins of México and Zacapu, Quatern. Int., 43, , Lumley, S. H. and Switsur, R.: Late Quaternary of the Taitao Peninsula, southern Chile, J. Quaternary Sci., 8, , Maberly, D. J.: The Plant Book: A portable dictionary of the higher plants, Cambridge University Press, 707 pp., Mancini, M. V.: Recent pollen spectra from forest and steppe of South Argentina: a comparison with vegetation and climate data, Rev. Palaeobot. Palyno., 77, , Marchant, R. A., Almeida, L., Behling, H., Berrio, J. C., Bush, M., Cleef, A., Duivenvoorden, J., Kappelle, M., De Oliveira, P. E., Hooghiemstra, H., Teixeira de Oliveira-Filho, A., Ledru, M.- P., Lozano-García, M. S., Ludlow-Wiechers, B., Mancini, V., Paez, M., Markgraf, V., Prieto, A., Rangel, O., and Salgado-Labouriau, M. L.: Distribution and ecology of parent taxa of pollen lodged within the Latin American Pollen Database, Rev. Palaeobot. Palyno., 121, 1 7, 02c. Marchant, R. A., Behling, H., Berrio, J. C., Cleef, A., Duivenvoorden, J., van Geel, B, van der Hammen, T., Hooghiemstra, H., Kuhry, P., Melief, B. M., van Reenen, G., and Wille, M.: Late Holocene Colombian vegetation dynamics, Quaternary Sci. Rev.,, , 01a. Marchant, R. A., Behling, H., Berrio, J. C., Cleef, A., Duivenvoorden, J., van Geel, B., van der Hammen, T., Hooghiemstra, H., Kuhry, P., Melief, B. M., van Reenen, G., and Wille, M.: A reconstruction of Colombian biomes derived from modern pollen data along an altitude gradient, Rev. Palaeobot. Palyno., 117, 79 92, 01b. Marchant, R. A., Behling, H., Berrio, J. C., Cleef, A., Duivenvoorden, J., van Geel, B., van der Hammen, T., Hooghiemstra, H., Kuhry, P., Melief, B. M., van Reenen, G., and Wille, M.: Colombian vegetation derived from pollen data at 0, 3000, 6000, 9000, , 000 and radiocarbon years before present, J. Quaternary Sci., 17, , 02a. Marchant, R. A., Behling, H., Berrío, J. C., Cleef, A., Duivenvoorden, J., van Geel, B., van der Hammen, T., Hooghiemstra, H., Kuhry, P., Melief, B. M., van Reenen, G., and Wille, M.: Mid to Late Holocene vegetation disturbance in Colombia a regional reconstruction, Antiquity, 78, , 04b. Marchant, R. A., Berrío, J. C., Behling, H., Boom, A., and Hooghiemstra, H.: Colombian dry moist forest transitions in the Llanos Orientales a comparison of model and pollen-based biome reconstructions, Palaeogeogr. Palaeocl., 234, 28 44, 06. Marchant, R. A., Boom, A., Behling, H., Berrío, J. C., van Geel, B., van der Hammen, T., Hooghiemstra, H., Kuhry, P., and Wille, M.: Colombian vegetation at the Late Glacial Maximum a comparison of model and pollen-based biome reconstructions, J. Quaternary Sci., 19, , 04 Marchant, R. A. and Hooghiemstra, H.: Rapid environmental change in tropical Africa and Latin America about 4000 years before present: a review, Earth-Sci. Rev., 66, , 04. Marchant, R. A., Boom, A., and Hooghiemstra, H.: Pollen-based biome and δ 13 C reconstruc- tions for the past years from the Funza II core, Colombia ecosystem relationships to late Quaternary CO 2 change as detected in the Vostok ice core record, Palaeogeogr. Palaeocl., 177, 29 4, 02b. Marchant, R. A. and Hooghiemstra, H.: Letter to the Editor Climate of East Africa C Yr BP as inferred from pollen data, By Odile Peyron, Dominique Jolly, Raymonde Bonnefille, Annie Vincens and Joël Guiot, Quaternary Res., 6, , 01b. Marchant, R. A. and Taylor, D. M.: Numerical analysis of modern pollen spectra and in situ montane forest implications for the interpretation of fossil pollen sequences from tropical Africa, New Phytol., 146, 0, 00. Markgraf, V.: Late and postglacial vegetational and paleoclimatic changes in sub-antarctic, temperate, and arid environments in Argentina, Palynology, 7, 43 2, Markgraf, V.: Late Pleistocene and Holocene vegetation history of temperate Argentina: Lago Morenito, Bariloche, Dissertationes Botanicae, 72, 23 4, Markgraf, V.: Late Pleistocene faunal extinction in southern Patagonia, Science, 228, , 198a. Markgraf, V.: Paleoenvironmental history of the last 000 years in northwestern Argentina, Zentralblatt für Geologie und Paläontolgie, 11/12, , 198b. 418

27 30 Markgraf, V.: Paleoenvironmental changes at the northern limit of the sub-antarctic Nothofagus forest, latitude 37S, Argentina, Quaternary Res., 28, , Markgraf, V.: Fells cave: years of changes in paleoenvironments, fauna, and human occupation, in: Travels in Archaeology in South Chile, edited by: Hyslop, J., University of Iowa, Iowa City, USA, 278 pp., Markgraf, V.: Late Pleistocene/Holocene paleoclimates from sub-antarctic latitudes, Antarct. J. US, 24, 1 2, Markgraf, V.: Palaeoclimates in Central and South America since BP based on pollen and lake-level records, Quaternary Sci. Rev., 8, 1 24, Markgraf, V.: Climatic history of Central and Southern America since yr BP, in: Global Climates since the Last Glacial Maximum, edited by: Wright, H. E., Kutzbach, J. E., Webb III, T., Ruddiman, W. F., Street-Perrot, F. A., and Bartlein, P. J., University of Minnesota Press, USA, 68 pp., Markgraf, V.: Younger Dryas in southern South America?, Boreas,, 63 69, Markgraf, V.: Paleoenvironments and paleoclimates in Tierra del Fuego and southernmost Patagonia, South America, Palaeogeogr. Palaeocl., 2, 3 68, Markgraf, V.: Past climates of South America, in: Climates of the Southern Continents: Present, Past and Future, edited by: Hobbs, E., Lindesay, J. A., and Bridgman, H. A., Wiley, Chichester, 342 pp., Markgraf, V., Baumgartner, T. R., Bradbury, J. P., Diaz, H. F., Dunbar, R. B., Luckman, B. H., Seltzer, G. O., Swetman, T. W., and Villalba, R.: Paleoclimate reconstruction along the Pole- Equator-Pole transect of the Americas (PEP 1), Quaternary Sci. Rev., 19, 1 140, 00. Markgraf, V., Bradbury, J. P., and Fernández, P.: Bajada de Rahue, Province Neuquen, Argentina: an interstadial deposit in northern Patagonia, Palaeogeogr. Palaeocl., 6, 1 8, Martin, L., Fournier, M., Mourguiart, P., Sifeddine, A., and Turcq, B.: Southern Oscillation signal in South American palaeoclimatic data of the last 7000 years, Quatern. Res., 39, , Martin, L., Bertaux, J., Correge, T., Ledru, M.-P., Mourguiart, P., Sifeddine, A., Soubiès, F., Wirrmann, D., Suguio, K., and Turcq, B.: Astronomical forcing of contrasting rainfall changes in tropical South America between and 8800 cal yr BP, Quatern. Res., 47, , Mayle, F. E., Burbridge, R., and Killeen, T. J.: Millennial-scale dynamics of southern Amazonian rain forest, Science, 290, , 00. Melief, A. B. M.: Late Quaternary palaeoecology of the Parque National los Nevados Cordillera Central, and Sumapaz Cordillera Oriental areas, Colombia, Ph.D. thesis, University of Amsterdam, Amsterdam, The Netherlands, 162 pp., 198. Mercer, J. H. and Ager, T. A.: Glacial and floral changes in southern Argentina since years ago, National Geographic Science Reports,, , Moreno, P. I.: Vegetation and climate near Lago Llanquihue in the Chilean Lake District between 0 and C yr BP, J. Quaternary Sci., 12, 48 00, Metclafe, S. E., O Hara, S. L., Caballero, M., and Davies, S. J.: Records of late Pleistocene- Holocene climate change in México a review, Quaternary Sci. Rev., 19, , 00. Monsalve, J. G.: A pollen core from the Hacienda Lusitania, Pro Calima, 4, 40 44, 198. Mourguiart, P. H., Argollo, J., and Wirrmann, D.: Evoucioun del lago Titicaca desde 000 nos BP, in: Climas cuaternarios en America del Sur, edited by: Argollo, J. and Mourguiart, P., Editions ORSTOM-La Paz, Bolivia, 7 171, 199. Noblet-Docoudré, N., Claussen, M., and Prentice, C.: Mid-Holocene greening of the Sahara: first results of the GAIM 6000 year BP experiment with two asynchronously coupled atmosphere/biome models, Clim. Dynam., 16, , 00. Northrop, L. A. and Horn, S. P.: Precolombian agriculture and forest disturbance in Costa Rica: palaeoecological evidence from two lowland rain forest lakes, Holocene, 6, , Ortega-Guerroro, B.: Paleomagnestismo, magnetoestrtifia y paleoecologia del Cuaternario tardio en el Lago de Chalco, Cuenca de México, Ph.D. thesis, National University of México, 8 pp., Páez, M. M. and Prieto, A. R.: Paleoenviornental reconstruction by pollen analysis from loess sequences from the Southeast of Buenos Aires (Argentina), Quatern. Int., 17, 139 3, Pendall, E., Markgraf, V., White, J. W. C., and Dreier, M.: Multiproxy record of late Pleistocene- Holocene climate and vegetation changes from a peat bog in Patagonia, Quaternary Res.,, , 01. Peng, C. H., Guiot, J., and van Campo, E.: Estimating changes in terrestrial vegetation and carbon storage: using palaeoecological data and models, Quaternary Sci. Rev., 17, ,

28 30 Pereira, B. A. S., de-mendonça, R. C., Filgueiras, T. S., de-paula, J. E., and Heringer, E. P.: Levantamento florístico da área deproteção ambiental (APA) da bacia do Rio São Bartolomeu, Distrito Federal, Anais XXXVI Congresso Brazileiro de Botânica, Socied, Bot. Brazil, 198, edited by: IBAMA, Curitiba, 1, Peyron, O., Jolly, D., Bonnefille, R., Vincens, A., and Guiot, J.: Climate of East Africa C yr BP as inferred from pollen data, Quaternary Res., 4, 90 1, 00. Pickett, E. J., Harrison, Sandy P., Hope, G., Harle, K., Dodson, J. R., Kershaw, A. P., Prentice, I., Colin, Backhouse, J., Colhoun, E. A., D Costa, D., Flenley, J., Grindrod, J., Haberle, S., Hassell, C., Kenyon, C., Macphail, M., Martin, H., Martin, A. C., McKenzie, M., Newsome, J. C., Penny, D., Powell, J., Raine, J. I., Southern, W., and Stevenson, J.: Pollen-based reconstructions of biome distributions for Australia, Southeast Asia and the Pacific (SEAPAC region) at 0, 6000 and C 14 yr BP, J. Biogeogr., 31, , 04. Piperno, D. R., Bush, M., and Colinvaux, P. A.: Palaeoenvironments and human occupation in Late Glacial Panama, Quaternary Res., 33, 8 116, Piperno, D. R., Bush, M., and Colinvaux, P. A.: Palaeoecological perspectives on human adap- tion in Central Panama. I: The Pleistocene, Geoarchaeology, 6, 1 226, 1991a. Piperno, D. R., Bush, M., and Colinvaux, P. A.: Palaeoecological perspectives on human adaption in Central Panama. II: The Holocene, Geoarchaeology, 6, 227 0, 1991b. Piperno, D. R., Ranere, A. J., Holst, I., and Hansell, P.: Starch grains reveal early root crop horticulture in the Panamanian tropical forest, Nature, 407, , 00. Pires, J. M. and Prance, G. T.: The vegetation types of the Brazilian Amazon, in: Key Environments Amazonia, edited by: Prance, G. T. and Lovejoy, T., Pergamon Press, 9 14, 198. Prance, G. T.: The changing forest, in: Key Environments Amazonia, edited by: Prance, G. T. and Lovejoy, T., Pergamon Press, 14 16, 198. Prentice, I. C., Cramer, W., Harrison, S. P., Leemans, R., Monserud, R. A., and Solomon, A. M.: A global biome model based on plant physiology and dominance, soil properties and climate, J. Biogeogr., 19, , Prentice, I. C., Sykes, M. T., Lautenschlager, M., Harrison, S. P., Denissenko, O., and Bartlein, P.: Modelling global vegetation patterns and terrestrial carbon storage at the last glacial maximum, Global Ecol. Biogeogr., 3, 67 76, Prentice, I. C., Guiot, J., Huntley, B., Jolly D., and Cheddadi, R.: Reconstructing biomes from palaeoecological data: a general method and its application to European pollen data at and 6 ka, Clim. Dynam., 12, , 1996a. Prentice, I. C., Harrison, S. P., Jolly D., and Guiot, J.: The climate and biomes of Europe at 6000 yr BP: comparison of model simulations and pollen-based reconstructions, Quaternary Sci. Rev., 17, , 1996b. Prentice, I. C. and Webb III, T.: BIOME 6000: reconstructing global mid-holocene vegetation patterns from palaeoecological records, J. Biogeogr.,, 997 0, Prentice, I. C., Jolly, D., and BIOME 6000 Participants: Mid-Holocene and glacial-maximum vegetation: geography of the northern continents and Africa, J. Biogeogr., 27, 07 19, 00. Prieto, A. R.: Late Quaternary vegetation and climatic changes in the pampa grassland of Argentina, Quaternary Res., 4, 73 88, Prieto, A. R. and Páez, M. M.: Pollen analysis of discontinuous stratigraphical sequences: Holocene at Cerro La China locality (Buenos Aires, Argentina), Quateranry of South America and Antartica Peninsula, 7, , Rodgers, J. C. and Horn, S. P.: Modern pollen spectra from Costa Rica, Palaeogeogr. Palaeocl., 124, 3 71, Rull, V.: A palynological record of a secondary succession after fire in Gran Sabana, Venezuela, J. Quaternary Sci. 14, 137 2, Rull, V., Salgado-Labouriau, M. L., Schubert, C., and Valastro Jr., S.: Late Holocene temperature depression in the Venezuelan Andes: Palynological evidence, Palaeogeogr. Palaeocl., 60, 9 121, Rzedowski, V.: Vegetacion de México. Editorial Lémusa, México, 432 pp., Salami, M.: Additional information on the findings in the Mylodo cave at Ultima Esperanza, Acta Geogr., 14, , 19. Salgado-Labouriau, M. L.: Sequence of colonisation by plants in the Venezuelan Andes after the Pleistocene glaciation, Journal of palynology, 23 24, 189 4, Salgado-Labouriau, M. L.: Late Quaternary palaeoclimate in the savannas of South America, J. Quaternary Sci., 12, , Salgado-Labouriau, M. L., Barberi, M., Ferraz-Vicentini, K. R., and Parizzi, M. G.: A dry climatic event during the late Quaternary of tropical Brazil, Rev. Palaeobot. Palyno., 99, 1 129, Salgado-Labouriau, M., Ferraz-Vicentini, K. R., and Parizzi, M. G.: A dry climatic event during the late Quaternary of tropical Brazil, Rev. Palaeobot. Palyno., 99, 129,

29 30 Saporito, M. S.: Chemical and mineral studies of a core from Lake Patzcuaro, México, MSc. thesis, University of Minnesota, Minneapolis, Minnesota, USA, 482 pp., 197. Sarmiento, G.: The dry plant formations of South America and their floristic connections, J. Biogeogr., 2, 233 1, 197. Schäbitz, F.: Untersuchungen zum aktuellen Pollenniederschlag und zur Holozänen Klima- und Vegetationsentwicklung in den Anden Nord-Neuquéns, Argentinien, Bamberger Geographische Schriften, 8, 1 131, Schäbitz, F.: Holocene vegetation and climate in southern Santa Cruz, Argentina, Bamberger Geographische Schriften, 11, , Schäbitz, F.: Holocene climatic variations in northern Patagonia, Argentina, Palaeogeogr. Palaeocl., 9, , Schreve-Brinkman, E. J.: A palynological study of the upper Quaternary sequence in the El Abra corridor and rock shelters Colombia, Palaeogeogr. Palaeocl.,, 1 9, Schmithüsen, J.: Atlas zur Biogeographie, Bibliographisches Institut, Mannheim/Wien/Zürich, 80 pp., Schofield, K.: Plants of the Galapagos Islands. Universe Books, 364 pp., Seibert, P.: Farbatlas Südamerika Landschaften und Vegetation, Verlag Eugen Ulmer, Mannheim, 288, Seltzer, G. O.: Recent glacial history and palaeoclimate of the Peruvian-Bolivian Andes, Quaternary Sci. Rev., 9, 137 2, Servant, M., Maley, J., Turcq, B., Absy, M., Brenac, P. Furnier, M., and Ledru, M.-P.: Tropical Forest changes during the late Quaternary in African and South American lowlands, Global Planet. Change, 7, 40, Sifeddine, A., Martin, L., Turcq, B., Volkermer-Ribero, C., Soubies, F., Cordeiro, R. C., and Suguio, K.: Variations in the Amazonian rain forest environment: a sedimentological record covering years, Palaeogeogr. Palaeocl., 168, , 01. Steege, H. T.: The use of forest inventory data for a National Protected Area Strategy in Guyana, Biodivers. Conserv., 7, , Takahara, H., Sugita, S., Harrison, S. P., Miyoshi, N., Morita, Y., and Uchiyama, T.: Pollen- Based Reconstruction of Japanese Biomes at 0, 6000 and yr BP, J. Biogeogr., 27, , 01. Tarasov, P. E., Webb III, T., Andreev, A. A., Afanas eva, N. B., Berezina, N. A., Bezusko, L. G., Blyakharchuk, T. A., Bolikhovskaya, N. S., Chernavskaya, M. M., Chernova, G. M., Doro feyuk, N. I., Dirksen, V. G., Elina, G. A., Filimonova, L. V., Glebov, F. Z., Guiot, J., Gunova, V. S., Harrison, S. P., Jolly, D., Khomutova, V. I., Kvavadze, E. V., Osipova, I. M., Panova, N. K., Prentice, I. C., Saarse, L., Sevastyanov, C. S., Volkova, V. S., and Zernitskaya, V. P.: Present day and mid-holocene Biomes Reconstructed from Pollen and Plant Macrofossil Data from the Former Soviet Union and Mongolia, J. Biogeogr.,, 29 3, 1998a. Tarasov, P. E., Cheddadi, R. Guiot, J., Sytze, B., Peyron, O., Belmonte, J., Ruiz-Sanchez, V, Saadi, F., and Brewer, S.: A method to determine warm and cool steppe biomes from pollen data; application to the Mediterranean and Kazakhstan regions, J. Quaternary Sci., 13, , 1998b. Texier, D. de Noblet, N., Harrison, S. P., Haxeltine, A., Joussaume, S., Jolly, D., Laarif, F., Prentice, I. C., and Tarasov, P. E.: Quantifying the role of biosphere-atmosphere feedbacks in climate change: a coupled model simulation for 6000 yr BP and comparison with palaeodata for Northern Eurasia and northern Africa, Clim. Dynam., 13, , Thompson, L. G., Mosely-Thompson, E., Davis, E. M., Lin, P. E., Henderson, K. A., Cole-Dai, B., and Liu, K.: Late glacial stage and Holocene tropical ice core records from Huascaren, Peru, Science, 269, 46 0, 199. Thompson, R. S. and Anderson, K. H.: Biomes of Western North America at , 6000 and 0 14 C yr BP, Reconstructed from Pollen and Packrat Midden Data, J. Biogeogr., 27, 6 664, 00. Thouret, J., van der Hammen, T., Salomons, B., and Juvigne, E.: Late Quaternary glacial studies in the Cordillera Central, Colombia, based on glacial geomorphology, tephra-soil stratigraphy, palynology and radiocarbon dating, J. Quaternary Sci., 12, , Thomas, M. F.: Late Quaternary environmental changes and the alluvial record in humid tropical environments, Quaternary Int., 72, 23 36, 00. Tuomisto, H. and Ruokolainen, K.: The role of ecological knowledge in explaining biogeography, and biodiversity in Amazonia, Biodivers. Conserv., 6, , Valencio, A. A., Creer, K. M., Sinito, A. M., Mazzoni, M. M., Alonso, M. S., and Markgraf, V.: Palaeomagnetism, sedimentology, radiocarbon age determinations and palynology of the Llao area, southwestern Argentina (lat. 41 S, long W): paleolimnological aspects, Quaternary of South America and Antarctic Penninsular, 3, 9 148, 198. Van der Hammen, T.: Palinología de la region de Laguna de los Bobos, Hitoria de su clima, vegetacion y agricultura durante los últimos 000 anos, Revista Acad Colombiana Ciencias Exactas Fisicas Naturales, 11, ,

30 30 Van der Hammen, T.: A Palynological Study on the Quaternary of British Guyana, Leidse Geologische Mededelingen, 29, , Van der Hammen, T.: Changes in vegetation and climate in the Amazon Basin and surrounding areas during the Pleistocene, Geol. Mijnbouw, 1, , Van der Hammen, T.: The Pleistocene changes of vegetation and climate in tropical South America, J. Biogeogr., 1, 3 26, Van der Hammen, T. and Absy, M.: Amazonia during the last glacial, Palaeogeogr. Palaeocl., 9, , Van der Hammen, T., Barelds, T. J., de Jong, H., and de Veer, A. A.: Glacial sequence and environmental history in the Sierra Nevada del Cocuy Colombia, Palaeogeogr. Palaeocl., 32, , Van der Hammen, T. and González, E.: Upper Pleistocene and Holocene climate and vegetation of the Saban de Bogotá, Leidse Geologische Mededelingen,, 261 3, Van der Hammen, T. and González, E.: A Late glacial and Holocene pollen diagram from Cienaga del Vistador Dep. Boyaca, Colombia, Leidse Geologische Mededelingen, 32, 193 1, 196. Van Geel, B. and Van der Hammen, T.: Upper Quaternary vegetation and climatic sequence of the Fuquene area Eastern Cordillera, Colombia, Palaeogeog. Palaeocl., 14, 9 92, Valdes, P.: South American palaeoclimate model simulations: how reliable are the models, J. Quaternary. Sci.,, , 00. Veblen, T. T., Burns, B. R., Kitzberger, T., Lara, A., and Villalba, R.: The Ecology of the Conifers of southern South America, in: Ecology of southern Conifers, edited by: Enright, N. J. and Hill, R. S., Melbourne University Press, 384 pp., 199. Vernet, J.-L., Wengler, L., Solari, M.-E., Ceccantini, G., Fournier, M., Fournier, M., Ledru, M.- P., and Soubies, F.: Feux, climats et végétations au Brésil central durant l Holocène: les donnees dún profil de sol a charbons de bois (Salitre, Minas Gerais), CRAS, Paris, 319, , Villagran, C.: Expansion of Megallanic Moorland during the Late Pleistocene: palynological evidence from Northern Isla de Chiloé, Chile, Quaternary Res., 30, , Watts, W. A. and Bradbury, J. P.: Palaeoecological studies on Lake Patzcuaro on the wets- central Mexican Plateau and at Chalco in the Basin of México, Quaternary Res., 17, 6 70, Webb III, T.: A global paleoclimatic data base for 6000 yr BP, US Department of Energy, Washington, DOE/EV/097 6, 198. Wijmstra, T. A. and van der Hammen, T.: Palynological data on the history of tropical savannas in northern South America, Leides Geologische Mededelingen, 38, 71 90, Wille, M., Hooghiemstra, H., Behling, H., Van der Borg, K., and Negret, A. J.: Environmental change in the Colombian subandean forest belt from 8 pollen records: the last 0 kyr, Veg. Hist. Archaeobot.,, 61 77, 01. Wille, M., Negret, J. A., and Hooghiemstra H.: Paleoenvironmental history of the Popayan area since yr BP at Timbio, Southern Colombia, Rev. Palaeobot. Palyno., 9, 4 63, 00. Wijmstra, T. A. and van der Hammen, T.: Palynological data on the history of tropical savannas in northern South America, Leidse Geologische Mededelingen, 38, 71 90, Wijmstra, T. A.: The Palynology of the Guiana coastal Basin, University of Amsterdam, 72 pp, Whitmore, T. C.: Quaternary History of tropical America, in: Biogeography and Quaternary History in Tropical America, edited by: Haffer, J., Clarendon Press, Oxford, 1 8, Williams, J. W., Summer, R. S., and Webb III, T.: Applying Plant Functional types to construct biome maps from eastern North American Pollen Data: Comparisons with model results, Quaternary Sci. Rev., 17, , Williams, J. W., Webb III, T., Richard, P. J. H., and Newby, P.: Late Quaternary Biomes of Canada and the Eastern United States, J. Biogeogr., 27, 23 38, 00. Wingenroth, M. and Suarez, J.: Flores de los Andes Alta montaña de Mendoza, IANIGLA (Instituto Argentino de Nivolgia y Glacologia, Argentina, 139 pp., Witte, H. J. L.: Present and past vegetation and climate in the North Andes (Cordillera Central, Colombia); a quantitative approach, Ph.D. thesis, University of Amsterdam, The Netherlands, 269 pp., Woodward, F. I.: Climate and plant distribution, Cambridge University Press, Cambridge, 188 pp., Yu, G. and Harrison, S. P.: Lake status records from Europe: data base ducumentation, NOAA, Boulder, Colorado, NOAA Paleoclimatology Publication Series, report, 3, 1 41, 199. Yu, G., Prentice, I. C., Harrison, S. P., and Sun, X.: Pollen-based biome reconstruction for China at 0 and 6000 years, J. Biogeogr.,, 69,

31 Yu, G., Xudong, C., Jian, N., Cheddadi, R., Guiot, J., Huiyou, H., Harrison, S. P., Ci-xuan, H., Jolly, D., Manhong, K., Zhaochen, K., Shengfeng, L., Wen-yi, L., Liew, P. M., Gunagxu, L., Jinling, L., Liu, K-B., Prentice, I. C., Guoyu, R., Changqing, S., Sugita, S., Xiangjun, S., Lingyu, T., Van Campo, E., Yumei, X., Qinghai, X., Shun, Y., Xiangdong Y., and Zhuo, Z.: Palaeovegetation of China: a pollen data-based synthesis for the mid-holocene and last glacial maximum, J. Biogeogr., 27, , 01. Xelhuantzi Lopez, M. S.: Determinacion palinologicá del paléoambiente holocenico en la parte norte del estado de Michoancán, Boletn Societe Botanica Mexicao, 4, 1 26, Xelhuantzi Lopez, M. S.: Palynologie et paléoenvironment du bassin de Zacapu, Michoancán, Mexique, depuis 8000 ans, Geofisica Internacional, 34, , Table 1. Characteristics of the sites from Latin America: specifically detailing location, potential vegetation around the sites, sample type, age range of sediments, number of C14 dates, data type, principle analysts and associated references. Dating control (DC) codes are based on the COHMAP dating control scheme (Webb, 198; Yu and Harrison, 199). For sites with a continuous sedimentation (indicated by C after the numerical code), the dating control is based on bracketing dates as follows. 1: both dates within 00 years of the selected interval, 2: one date within 00 years the other within 4000 years, 3: both dates within 4000 years, 4: one date within 4000 years one date within 6000 years, : both dates within 6000 years, 6: one date within 6000 years the other within 8000 years, 7: bracketing dates more than 8000 years from the selected interval. For sites with discontinuos sedimentation (indicated by D after the numerical code), the dating control is based on single dates 1: indicated a date within 0 years of the selected interval, 2: a date within 00 years, 3: a date within 70 years, 4: a date within 00 years, : a date within 00 years, 6: a date within 00 years, 7: a date of more than 00 years from the selected interval. Code Biome Definition Main locations Equivalent Floristic characteristics TRFO Tropical Closed canopy lowland Characterise much of Amazonian forest, Generally characterised by rainforest evergreen forests. the Amazon catchment. Can form tropical moist forest, plants with mesophyll leaf, Canopy broken by a relatively thin band along Atlantic rain forest, although some sclerophyllous emergent trees (>40 m). tropical coastal areas, e.g. Atlantic terra firme forest, plants are present, MTCO >18 C, rain forest of Brazil, Chocó Várzea, Gallery forest, often tree ferns and palms. α>00, pluvial forest of Colombia, Chocó pluvial forest frost-intolerant. maintained by high moisture derived from close proximity of oceanic influence. TSFO Tropical Relatively tall ( 30 m) Dominant to the north Marsh forests, A mix of mesophyllous and seasonal closed canopy forest with of Amazonian tropical savanna gallery forest, sclerophyllous taxa. forest occasionally tall (>40 m) rainforest, in central America Seasonal swamp forest The structure of the forest emergent trees. and formerly extensive with palms. is dependent on moisture Canopy opens in a mosaic in the interior of Brazil demand and length of dry season as deciduous elements prior to extensive clearance. this determines the amount loose leaves. Seasonally of deciduous taxa. Palms dry from 1 4 months. can be locally common. TDFO Tropical Relatively low ( m), Extensive in central Brazil where Andean xerophytic bush, Xeromorphic characteristic, dry occasionally tall ( m) it abuts tropical rainforest. Cerrado, Campo rupestres, dparticularly rought and fire tolerant. forest trees. Mixed forest, More fragmented in northwestern Campo cerrado ( campo is For example, microphyllous leaves, forming where the dry South America where a free draining more associated with thorns, deciduous leaves, thick bark, season leads to drought substrate leads to water-stress. grasslands). Cactus forest, stomata often present along lines. and plant water stress. Extends to mid altitudes, particularly Matorral, Deciduous xerophytic Drought adapted taxa are common, rwithin ain shadow areas. Extensive forest, Andean xerophytic bush, e.g. tree cacti (Opuntia in western Central America and Espinar, Restinga dune Tforests, and Jasminocerus) with Mexico. Present on the Galápagos horn forest, Chaco. dense undergrowth of shrubs Islands, extensive in Chile, central and herbs. South America. Associated with the Andes, particularly within rain shadow areas. Extends into dry areas of Central America such as the Yucatan peninsular. 428

32 Table 1. Continued. Code Biome Definition Main locations Equivalent Floristic characteristics WTRF Warm Evergreen closed forest, Extending along the Andes Lower montane forest, A mix of mesophyllous temperate of relatively low stature (< m) at mid to low elevations (00 00 m). moist lower montane forest, and sclerophyllous taxa rainforest with tall emergent trees (> m). Present at slightly lower elevations submontane forest, constrained by altitude Not tolerant of freezing. in eastern Brazilian highlands (<00 m). subandean forest, and length of dry A transitional forest type The similarity with the Andean Araucaria-dominated forest season. Palms and tree between lowland and higher flora indicates these areas have also with Podocarpus. ferns can be locally common. altitude forms (00-00 m). been connected in the past WEFO Warm Evergreen semi-closed Present within a relatively Andean forest, A mix of mesophyllous temperate forest with tall restricted range transitional Andean forest, and sclerophyllous taxa. evergreen emergent trees (>30 m). and along the Andes, upper Andean forest Tree ferns can be broadleaf Not tolerant of freezing. particularly present locally common. forest from m. CTRF Cool Medium height (< m) Predominant along western Patagonian rain forest, A mix of mesophyllous temperate closed canopy forest coast of southern temperate rain forest, and sclerophyllous taxa. rainforest with a dense under-story. South America extending valdivian rain forest, The structure can be Can tolerate freezing. to Patagonian steppe. magallenic rain forest quite variable depending Also present along the Andes on location from at mid to high altitudes. dense forest to scrubby heath. WAMF Warm Medium height (< m) Mid to high altitudes Pinus and Mixed evergreen forest temperate open canopy with open of north Central Quercus-dominated dominated by sclerophyllous mixed under-story. Drought America, in particular forest. taxa that require forest tolerant, semi fire-tolerant. Mexico warm for bud-burst. COMI Cool Short stature woodlands (< m) High Andean shrub/dwarf Upper montane forests, Predominantly evergreen mixed open canopy, open under-story tree forests, present close high Andean forest, taxa with physiological forest forest. Frost tolerant. to the forest line cloud forest adaptation to night frost, e.g. retaining old leaves for insulation. CGSS Cool Common above the forest Present only at Puna, Heath, Poaceae-dominated cool grasslands line of the Andes, the highest altitudes Cushion heath grasslands with occasional dominated by tussock of the Andes cushion plants grasses and cusion plants. STEP Steppe Dominated by grasses, Extensive in eastern Argentina, Steppe grasslands, Grasses and chenopods occasional shrubs and present in lowland Central America Campo limpo, forming low altitude steppe herbs. Profuse and northeast Brazil Pampa warm grasslands flowering during the wet season DESE Desert Open semi-arid Coastal Peru and Chile and Coastal desert Occurrence of CAM-plants, to arid vegetation western Mexico, the former cacti and succulents area due to rain-shadow from the Andes CGSH Cool Tropic-alpine environments, Present from extreme southern Páramo, Subpáramo, Poaceae-dominated cool grass common above the forest line South America, on Tierre Magallenic moorland, grasslands with numerous shrublands of the Andes. A mixture del Fuego above the forest line Paramillo, Vegas tussock forming grass. of tussock grasses and of the Andes ( m). Also present are shrubs, cold-adapted shrubs. e.g. Empetrum, Espeletia and Puya. 429 Table 2. Range of plant functional types identified within the Latin American region giving bioclimatic range and physiological adaptation. Code Plant functional type Bioclimatic range and plant physiological adaption g Graminoid Ecologically broad category that occurs in a number of biomes, a highly adaptive PFT with a ubiquitous distribution and little diagnostic value. man Mangrove Constituent of lowland tropical vegetation, control on distribution is mainly hydrological tx Tree fern Can be locally dominant. Occupying a broad bioclimatic range, recorded in a range of moist environments from lowland to montane, particularly common in temperate areas. Te 1 Tropical mesic drought-deciduous broad-leaved tree MTCO>. C, α>0.7, short dry season (1 month), GDD>000 Te 2 Tropical xeric drought-deciduous broad-leaved tree MTCO>. C, α>0.6 08, longer dry season (2 4 months), GDD>000, withstands longer dry season by shedding leaves Tr 1 Tropical evergreen broad-leaved tree MTCO>. C, α>0.9, GDD>000, present in wettest tropical rain forest. Tr 2 Tropical mesic evergreen broad-leaved tree MTCO>. C, α , GDD>000, present in range of tropical seasonal forest types ctc Warm temperate evergreen needle-leaved tree MTCO C C, α>0.7, GDD>400, common in the Brazilian highlands ctc 1 Cool temperate evergreen needle-leaved tree MTCO C C, α , GDD>900, common along the western coast of southern South America ctc 2 Cold evergreen needle-leaved tree MTCO C C, α>0.6, GDD>00, common along the western coast of southern South America ec Eurythermic conifer MTCO> C, α , GDD>000, common within dry forest of South America and Mexico txts Drought-tolerant small-leaved low or high shrub MTCO> C, α , GDD>000, woody shrubs common in dry forest ds Desert shrubs MTCO> C, α , GDD>000, woody shrub and cacti in Mexico and coastal Peru df Eurythermic drought-adapted forb MTCO> C, α , GDD>000, woody shrub and cacti in Mexico and coastal Peru tf Tropical drought-intolerant forb MTCO>. C, α>0.6, GDD>000, frost intolerant tef Temperate drought-intolerant forb MTCO> C C, α>0.6, GDD>00, frost tolerant sf Eurythermic drought-tolerant forb MTCO C C, α , GDD , requires a seasonal moist environment af Arctic forb MTCO C 0 C, α , GDD<00, frost tolerant cp Rosette or cushion forb MTCO< C, α<0.2, GDD<00, specific growth form, frost tolerant. wte Warm temperate evergreen broad-leaved tree MTCO C C, α>0.6, GDD>3000, frost tolerant mesophyllous trees ts Temperate evergreen broad-leaved tree MTCO 0 C C, α>0.6, GDD>00, frost tolerant micro and mesophyllous trees ts 1 Temperate evergreen sclerophyll broad-leaved tree MTCO C C, α>0., GDD>00, sclerophyllous, usually evergreen wte 1 Temperate (spring-frost avoiding) cold-deciduous MTCO 0 C C, α>0.6, GDD>3000, winter deciduous, requires broad-leaved tree warm growing season. wte 4 Temperate (spring-frost tolerant) cold-deciduous MTCO C C, α 0., GDD>00, winter deciduous, requires warm broad-leaved tree growing season but this can be short. aa Arctic evergreen broad-leaved erect dwarf shrub MTCO C 0 C, α , GDD 00 00, frost tolerant 430

33 Table 3. Biomes identified within the Latin American region as portrayed in the vegetation map (Fig. 2) indicting a floristic description, the main location and equivalent floristic units found in a macro scale analysis of the Latin American vegetation. Site Country Long. Latitude Alt. Age Present Sample RC DC at DC at Data Analyst Site range biome type type publications yr BP yr BP Lake Åsa 3 Antarctica CGSH Soil Raw Björck, S. Björck et al. (1993) Salina Anzotegui Argentina STEP Playa 3D - Raw Schäbitz, F. Schäbitz (1994) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) La Misión Argentina STEP Mire 4 4C - Raw Markgraf, V. Markgraf (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Harberton Argentina STEP Mire 16 1C - Raw Markgraf, V. Markgraf (1989, 1991, 1993) Ruta 3.4 Argentina Modern STEP Soil Raw Schäbitz, F. Schäbitz (1994) Ruta 3.3 Argentina Modern STEP Soil Raw Schäbitz, F. Schäbitz (1994) Pedro Luro Argentina Modern STEP Soil Raw Schäbitz, F. Schäbitz (1994) Origone Argentina Modern STEP Soil Raw Schäbitz, F. Schäbitz (1994) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Espuma Argentina Modern TDFO Soil Raw Schäbitz, F. Schäbitz (1994) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Arroyo Sauce Chico Argentina Modern STEP Soil Raw Prieto, A.R. Prieto (1996) Gaviotas Argentina Modern TDFO Soil Raw Schäbitz, F. Schäbitz (1994) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Empalme Querandíes Argentina STEP Lake 8 2C - Raw Prieto, A. R. Prieto (1996) Ruta 0.19 Argentina Modern STEP Soil Raw Schäbitz, F. Schäbitz (1994) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) 431 Table 3. Continued. Site Country Long. Latitude Alt. Age Present Sample RC DC at DC at Data Analyst Site range biome type type publications yr BP yr BP Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Moreno Glacier Bog Argentina CTRF Mire 2 4C - Raw Ager, T. A. Mercer and Ager (1983) Patagonia Argentina Modern CTRF Soil Digi Mancini, M. V. Mancini (1993) Cerro La China Argentina STEP Soil 2 4C - Raw Prieto, A. R. Prieto and Páez (1989), Páez and Prieto (1993) LagoBsAs Argentina Modern CTRF Soil Raw Schäbitz, F. Schäbitz (1994) Pico Salam Argentina Modern STEP Soil Raw Schäbitz, F. Schäbitz (1994) AlercesNor Argentina Modern CTRF Soil Raw Schäbitz, F. Schäbitz (1994) Mallin Book Argentina CTRF Mire 9 1C - Raw Markgraf, V. Markgraf (1983) Primavera Argentina CTRF Midden Raw Markgraf, V. Markgraf et al. (in press ) Comallo Argentina Modern CGSH Soil Raw Schäbitz, F. Schäbitz (1994) Encantado Argentina CTRF Midden Raw Markgraf, V. Markgraf et al. ( in press ) Meseta Latorre 1 Argentina CGSH Mire 3 4C - Raw Schäbitz, F Schäbitz (1991) Meseta Latorre 2 Argentina CGSH Mire 1 6D - Raw Schäbitz, F. Schäbitz (1991) Cueva Haichol Argentina STEP Cave 2 1C - Raw Markgraf, V. Markgraf (1988) AustroEsqu Argentina Modern CTRF Soil Raw Schäbitz, F. Schäbitz (1994) Vaca Lauquen Argentina CTRF Mire 3 1C - Raw Markgraf, V. Markgraf (1987) Veranada Pelan Argentina CGSH Mire 3 3C - Raw Schäbitz, F. Schäbitz (1989) Salina 2 Argentina CGSH Mire 2 1C - Raw Markgraf, V. Markgraf (1983) Veranada Vulkanpickel Argentina CGSH Mire 1 7D - Raw Schäbitz, F. Schäbitz (1989) Salado Argentina CGSH Mire Raw Markgraf, V. Markgraf (1983) Aguilar Argentina CGSH Mire 3 2C - Raw Markgraf, V. Markgraf (198) Laguna Bella Vista Bolivia TSFO Lake 2C 1D Digi Mayle, F. Mayle et al. (00) Laguna Chaplin Bolivia TSFO Lake 14 2C 1D Digi Mayle, F. Mayle et al. (00) Wasa Mayu Bolivia COMI Lake 1 7D 7D Raw Graf, K. Graf (1992) Lake Huinãmimarca Bolivia CSGH Lake 48 1C 3C Digi Mourguiart, P. H. Mourguiart et al. (199), Argollo and Mourguiart (00) Cerro Calvario Bolivia CGSH Mire 4 1C - Raw Graf, K. Graf (1992) Amarete Bolivia CGSH Mire 2 D - Raw Graf, K. Graf (1992) Rio Kaluyo Bolivia CGSH Lake 3 3C - Raw Graf, K. Graf (1992) Sajama Bolivia CGSH Lake Raw Graf, K. Graf (1992) Cotapampa Bolivia CGSH Mire 2C - Raw Graf, K. Graf (1992) Cumre Unduavi Bolivia CGSH Mire 6 3C - Raw Graf, K. Graf (1992) Chacaltaya 1 Bolivia CGSH Mire 1 1C - Raw Graf, K. Graf (1992) Mt. Blanco Bolivia CGSH Lake 7 1C - Raw Graf, K. Graf (1992) Katantica Bolivia CGSH Mire 3 1C - Raw Graf, K. Graf (1992) Reserva Volta Velha Brazil Modern WTRF Trap Digi Behling, H. Behling et al. (1997) Lago Crispim Brazil TRFO Lake 4 1C - Raw Behling, H. Behling et al. (1997) ODP site 932 Brazil TRFO Fan 2 2C 1D Raw Behling, H. Haberle and Maslin (1999) Lagoa da Caço Brazil TDFO Lake 14 1C - Raw Behling, H. Ledru et al. (01) Poço Grande Brazil WTRF Section Raw Behling, H. Behling (00), Behling (1997a, b) 432

34 Table 3. Continued. Site Country Long. Latitude Alt. Age Present Sample RC DC at DC at Data Analyst Site range biome type type publications yr BP yr BP Lagoa da Curuça 2 Brazil TRFO Lake 4 2C - Raw Behling, H. Behling (00) Rio (unclear) Brazil Modern TDFO River Raw Behling, H. Behling et al. (00) Rio Jaguaribe II Brazil Modern TDFO River Raw Behling, H. Behling et al. (00) Rio Jaguaribe I Brazil Modern TDFO River Raw Behling, H. Behling et al. (00) Picos Brazil Modern TDFO Soil - 1D 1D Raw Harbele, S. Behling et al. (00) Campina Grande I Brazil Modern TSFO Soil Raw Behling, H. Behling et al. (00) Rio Mirim Brazil Modern TDFO River Raw Behling, H. Behling et al. (00) Mirim Brazil Modern TDFO Soil - 1D 7D Raw Ledru, M.-P. Behling et al. (00) Lagoa Grande Brazil Modern TDFO Lake Raw Behling, H. Behling et al. (00) Rio de Contas Brazil Modern TDFO River Raw Behling, H. Behling et al. (00) Rio Jequitinhonha Brazil Modern TDFO River Raw Behling, H. Behling et al. (00) Rio São Francisco Brazil Modern TDFO River Raw Behling, H. Behling et al. (00) Rio Protengi Brazil Modern TDFO River Raw Behling, H. Behling et al. (00) Lago Bolim Brazil Moden TDFO Lake Raw Behling, H. Behling et al. (00) Comprida Brazil TRFO Lake 1C - Digi Bush, M. Bush et al. (00) Geral Brazil TRFO Lake 2 1C - Digi Bush, M. Bush et al. (00) Carajas Brazil TSFO Lake 8 2C 6C Digi Absy, M. L. Absy et al. (1991) Crominia Brazil TDFO Palm 2C 1D Digi Salgado- Salgado-Labouriau swamp -Labouriau, M. L. et al. (1998) Atlantic Brazil Modern WTRF Trap Raw Behling, H. Behling et al. (1997) Aguads Emendadas Brazil TDFO Palm 6 7C 7D Digi Salgado- Salgado-Labouriau swamp -Labouriau, M. L. et al. (1998) Lagoa das Patas Brazil TFFO Lake 16 1D 1C Raw De Oliveira, P. E. De Oliveira (1992) La Pata Brazil TRFO Lake 12 2C 1D Digi Colinvaux, P. Colinvaux et al. (1996, 00) Cuiaba Brazil Modern TDFO Soil Raw Behling, H. Behling (in prep ) Lago do Pires Brazil TSFO Lake 7 1C - Raw Behling, H. Behling (1993), Behling (1997a, b) Rio São Francisco Brazil TDFO River 6 1C - Digi De Oliveira, P. E. De Oliveira et al. (1999) Saquinho Brazil TSFO Mire 6 1C - Digi De Oliveira, P. E. De Oliveira et al. (1999) Assis Brazil Modern TSFO Soil Raw Behling, H. Behling (in prep ) Bauru Brazil Modern TSFO Soil Raw Behling, H. Behling (in prep ) Lagoa Santa Brazil Modern WTRF River Digi Parizzi, M.G. Salgado-Labouriau et al. (1998) Brotas Brazil Modern WTRF Soil Raw Behling, H. Behling (in prep ) Botucatu Brazil Modern WTRF Soil Raw Behling, H. Behling (in prep ) Curcuab Brazil Modern WTRF Soil Raw Behling, H. Behling (in prep ) Katira Brazil TDFO Lake 4 7D 1D Digi van der Hammen, T. van der Hammen and Absy (1994) Rio da Curuá Brazil WTRF Lake Raw Behling, H. Behling et al. (1997) Colombo Brazil Modern TSFO Trap Raw Behling, H. Behling et al. (1997) Brasilia 1 Brazil Modern TDFO Soil Raw Behling, H. Behling ( in prep ) Salitre Brazil WTRF Lake 14 1C 7C Raw Ledru, M.-P. Ledru (1992, 1993), Ledru et al. (1994, 1996) Serra da Boa Vista Brazil WTRF Mire 4 2C - Raw Behling, H. Behling (1993), Behling (1997a, b) Serra Campos Gerais Brazil WTRF Mire 4 3C - Raw Behling, H. Behling (1997a) Serra do Rio Rastro Brazil WTRF Mire 3 2C - Raw Behling, H. Behling (1993), Behling (1997a, b) Morro da Igreja Brazil WTRF Mire Raw Behling, H. Behling (1993), Behling (1997a, b) 433 Table 3. Continued. Site Country Long. Latitude Alt. Age Present Sample RC DC at DC at Data Analyst Site range biome type type publications yr BP yr BP Morro de Itapeva Brazil WTRF Lake 9 4C - Raw Behling, H. Behling (1997b) Puerto del Hambre Chile CTRF Mire 4C 7D Digi Heusser, C. Heusser et al. (199) Laguna Lofel Chile CTRF Lake 2C 6D Digi Bennet, K. Bennet et al. (00) Laguna Stibnite Chile CTRF Lake 6 1C 4D Digi Lumley, S. Lumley and Switsur (1993) Puerto Eden Chile CTRF Mire 7 4C - Raw Markgraf, V. Ashworth and Markgraf (1989), Ashworth et al. (1991) Laguna Stibnite Chile CTRF Lake 1C 2D Digi Bennet, K. Bennet et al. (00) Laguna Six Minutes Chile CTRF Lake 4 4C 7D Digi Bennet, K. Bennet et al. (00) Laguna Lincoln Chile CTRF Lake 2C 7D Digi Bennet, K. Bennet et al. (00) Dichan Chile CTRF Mire 2C - Digi Heusser, C. Heusser et al. (199) Estero Huitanque Chile CTRF Mire 9 1C - Digi Heusser, C. Heusser et al. (199) Rano Raraku Bore 3 Chile TDFO Lake 4C 4C Raw Flenley, J. Flenley and King (1984), Flenley et al. (1991) Mayol Chile CTRF Mire 12 3C - Digi Heusser, C. Heusser et al. (199) Punta Arenas Chile CTRF Mire 4C 7D Digi Heusser, C. Heusser et al. (199) Torres del Paine Chile CTRF Lake 8 2C - Digi Heusser, C. Heusser et al. (199) Rano Kao Chile TDFO Lake Raw Flenley, J. Flenley and King (1984), Flenley et al. (1991) Puchilco Chile CTRF Mire 7 2C - Digi Heusser, C. Heusser et al. (199) Puerto Octay PM13 Chile CTRF Mire 16 4C 1C Raw Moreno, P. I. Moreno (1994) Chepu Chile Modern CTRF Mire Raw Moar, N. T. Godley and Moar (1973) La Esperanza Chile CTRF Mire Raw Graf, K. Graf (1992) Rano Aroui Chile TDFO Lake 11 2C 6C Raw Flenley, J. Flenley and King (1984) Caunahue Chile CTRF Section 9 2C - Raw Markgraf, V. Markgraf (1991) San Pedro Chile Modern CTRF Mire Raw Moar, N. T. Godley and Moar (1973) Tumbre 2 Chile CGSH Lake 3 2C - Raw Graf, K. Graf (1992) Aguas Calientas Chile CGSH Mire 1 7D - Raw Graf, K. Graf (1992) Ajata Chile CGSH Mire Raw Graf, K. Graf (1992) Boca de Lopez Colombia TRFO Coastal - - Raw van der Hammen, T. Behling, Berrio and Hooghiemstra (1999) Jotaordó Colombia TRFO Lake Raw Berrio, J. C. B. Berrio, Behling and Hooghiemstra (00) El Caimito Colombia TRFO Lake Raw Wille, M. Wille et al. (1999) Monica-1 Colombia TRFO Lake 3 2C - Raw Behling, H. Behling, Berrio and Hooghiemstra (1999) Mariñame-II Colombia TRFO Lake 1C - Raw Behling, H. Behling, Berrio and Hooghiemstra (1999) Carimagua Colombia TDFO Lake 6 2C - Raw Behling, H. Behling and Hooghiemstra (1999) Sardinas Colombia TDFO Lake 6 2C - Raw Behling, H. Behling and Hooghiemstra (1998) El Piñal Colombia TDFO Lake 8 4C 2D Raw Behling, H. Behling and Hooghiemstra (1999) Piusbi Colombia TRFO Lake 3 1C - Raw Behling, H. Behling and Hooghiemstra (1999) Laguna Angel Colombia TDFO Lake 8 2C - Raw Behling, H. Behling and Hooghiemstra (1998) Lago Agua Sucia Colombia TDFO Lake 4 7D - Raw Wijmstra, T. A. Wijmstra and van der Hammen (1966) 434

35 Table 3. Continued. Site Country Long. Latitude Alt. Age Present Sample RC DC at DC at Data Analyst Site range biome type type publications yr BP yr BP Loma Linda Colombia TDFO Lake 8 1C - Raw Behling, H. Behling, Berrio and Hooghiemstra (1999) Pitalito Colombia WEFO Mire 6 6D - Raw Bakker, J. G. M. van der Hammen et al. (1980) Piagua Colombia WEFO Lake 7 7D - Raw Wille, M. Wille et al. (01), van der Hammen et al. (1980) Pantano de Genagra Colombia WEFO Mire 7 4C - Raw Behling, H. Behling, Negret and Hooghiemstra (1999), van der Hammen et al. (1980) Rio Timbio Colombia WEFO Lake 6 2C - Raw Wille, M. van der Hammen et al. (1980) Libano Colombia WEFO Soil 1 7D - Raw Salomons, J. B. van der Hammen et al. (1980) de Pedro Palo III Colombia COMI Lake 2 7D - Raw van der Hammen, T. van der Hammen (1974) Herrera Colombia COMI Lake 3 4D - Raw van Geel, B. van Geel and van der Hammen (1973) Ubaqué Colombia Modern WEFO Lake Raw Jean-Jarcob, K. Wille et al. (01), van der Hammen et al. (1980) Ciudad Universitaria X Colombia >3 000 COMI Lake 4 4C 7D Raw van der Hammen, T. van der Hammen and González (1960) El Abra II Colombia COMI Cave 1 7D - Raw Schreve-Brinkman, E. J. Schreve-Brinkman (1978) Fúquene II Colombia COMI Lake 2 7D 7C Raw van Geel, B. van Geel and van der Hammen (1973) Alsacia Colombia COMI Mire 3 6D - Raw Melief, A. B. M. Melief (198) Agua Blanca Colombia COMI Mire 2 6D 7D Raw Kuhry, P. Graf (1992), Kuhry (1988b), Kuhry et al. (1983) Cienaga del Visitador Colombia COMI Mire 2 7D - Raw van der Hammen, T. van der Hammen and González (196) La Guitarra Colombia COMI Mire 3 4C - Raw Melief, A. B. M. Melief (198) Ciega I Colombia COMI Lake Raw van der Hammen, T. van der Hammen et al. (1980) La Primavera Colombia CGSH Mire 6 1C - Raw Melief, A. B. M. Melief (198) de la América Colombia CGSH Mire 1 1D - Raw Kuhry, P. Kuhry (1988), van der Hammen and González (1960) Paramo Palacio Colombia CGSH Mire 4 D - Raw van der Hammen, T. van der Hammen and González (1960) Andabobos Colombia CGSH Mire 2 7D - Raw Melief, A. B. M. Melief (198) Paramo de Peña Negra Colombia CGSH Mire 2C - Raw Kuhry, P. Kuhry et al. (1983) Paramo de Laguna Verde Colombia CGSH Mire 2 4D - Raw Kuhry, P. Kuhry et al. (1983) El Bosque Colombia CGSH Mire Raw Melief, A. B. M. Kuhry (1988a), Melief (198) Turbera de Calostros Colombia Modern CGSH Soil Raw Salomons, J. B. van der Hammen et al. (1980) Bobos Colombia CGSH Lake 4 6D - Raw van der Hammen, T. van der Hammen (1962) El Gobernador Colombia CTRF Mire 2 2C - Raw Melief, A. B. M. Melief (198) Valle de Lagunillas Colombia CGSH Lake 8 7D - Raw van der Hammen, T. van der Hammen et al. (1980) La Rabona Colombia CGSH Mire 1 4D - Raw Melief, A. B. M. Melief (198) Greja Colombia CGSH Lake 2 3C - Raw van der Hammen, T. van der Hammen (1962) Corazón Partido Colombia Modern CGSH Mire Raw Melief, A. B. M. Melief (198) El Trinagulo Colombia Modern CGSH Mire Raw Melief, A. B. M. Melief (198) Santa Rosa1 Costa Rica Modern TRFO Soil Digi Horn, S. P. Rodgers and Horn (1996) Santa Rosa2 Costa Rica Modern TSFO Soil Digi Horn, S. P. Rodgers and Horn (1996) 43 Table 3. Continued. Site Country Long. Latitude Alt. Age Present Sample RC DC at DC at Data Analyst Site range biome type type publications yr BP yr BP Carara Biological Reserve Costa Rica Modern TSFO Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Tortuguero Costa Rica Modern TSFO Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Carara Costa Rica Modern TSFO Lake Digi Horn, S. P. Rodgers and Horn (1996) Cantarrana Swamp Costa Rica Modern TRFO Swamp Digi Horn, S. P. Rodgers and Horn (1996) Sendro Sedro Swamp Costa Rica Modern TRFO Swamp Digi Horn, S. P. Rodgers and Horn (1996) Laguna Palmita Costa Rica Modern TDFO Soil Digi Horn, S. P. Rodgers and Horn (1996) La Selva, Heredia Costa Rica Modern TSFO Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) La Pacifica, Guanacaste Costa Rica Modern TSFO Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Cataracta, Carara 1 Costa Rica Modern TSFO Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Cataracta, Carara 2 Costa Rica Modern TSFO Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Santa Rosa3 Costa Rica Modern TSFO Soil Digi Horn, S. P. Rodgers and Horn (1996) Santa Rosa4 Costa Rica Modern TSFO Soil Digi Horn, S. P. Rodgers and Horn (1996) Escondido Costa Rica Modern TSFO Lake Digi Horn, S. P. Rodgers and Horn (1996) Cafetal, Guanacaste Costa Rica Modern TSFO Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Laguna Río Cuarto Costa Rica Modern WTRF Lake Digi Horn, S. P. Rodgers and Horn (1996) Laguna Bonilla Costa Rica Modern TSFO Lake Digi Horn, S. P. Rodgers and Horn (1996) Laguna La Palma Costa Rica Modern WTRF Lake Digi Horn, S. P. Rodgers and Horn (1996) Laguna Cedeño Costa Rica Modern WTRF Lake Digi Horn, S. P Rodgers and Horn (1996) Brauillo Carillo, Heredia Costa Rica Modern WTRF Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Laguna González Costa Rica Modern WTRF Lake Digi Horn, S. P. Rodgers and Horn (1996) Laguna Congo Costa Rica Modern WTRF Lake Digi Horn, S. P. Rodgers and Horn (1996) Bosque Alegre Costa Rica Modern WTRF Lake Digi Horn, S. P. Rodgers and Horn (1996) Laguna Hule Costa Rica Modern WTRF Lake Digi Horn, S. P. Rodgers and Horn (1996) Laguna María Aguilar Costa Rica Modern WTRF Lake Digi Horn, S. P. Rodgers and Horn (1996) Volcán Cacao Costa Rica Modern WTRF Soil Digi Horn, S. P. Rodgers and Horn (1996) Monteverde, Heredia Costa Rica Modern WTRF Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) La Chonta Costa Rica CTRF Lake 3 2C 7D Digi Islebe, G. Hooghiemstra et al. (1992), Islebe and Hooghiemstra (1997), Islebe et al. (199a, b) Talamancas Costa Rica Modern CTRF Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Volcan Poas Costa Rica Modern CTRF Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Laguna Botos Costa Rica Modern CTRF Lake Digi Horn, S. P. Rodgers and Horn (1996) Tres de Junio Costa Rica Modern CTRF Bog Digi Horn, S. P. Rodgers and Horn (1996) Bog 68 Costa Rica Modern CTRF Bog Digi Horn, S. P. Rodgers and Horn (1996) Bog 70 Costa Rica Modern CTRF Bog Digi Horn, S. P. Rodgers and Horn (1996) Laguna Barva Costa Rica Modern CTRF Lake Digi Horn, S. P. Rodgers and Horn (1996) Quebrador Costa Rica Modern CTRF Lake Digi Horn, S. P. Rodgers and Horn (1996) Asuncion Costa Rica Modern CTRF Lake Digi Horn, S. P. Rodgers and Horn (1996) Lago de las Morrenas Costa Rica CTRF Lake 6 2C - Digi Horn, S. P. Horn (1993) Lago Chirripó Costa Rica Modern CTRF Lake Digi Horn, S. P. Rodgers and Horn (1996) 436

36 Table 3. Continued. Site Country Long. Latitude Alt. Age Present Sample RC DC at DC at Data Analyst Site range biome type type publications yr BP yr BP Limoncocha Ecuador TRFO Lake Digi Colinvaux, P. Colinvaux et al. (1988) Añangucocha Ecuador Modern TRFO Lake Digi Bush, M. Bush et al. (1990) Cuyabeno Ecuador Modern TRFO Lake Digi Bush, M. Bush et al. (1990) Lago Agrio Ecuador Modern TRFO Lake Digi Bush, M. Bush et al. (1990), Colinvaux et al. (1988) Santa Cecilia Ecuador Modern TRFO Lake Digi Bush, M. Bush et al. (1990) Lake Santa Cecilia Ecuador TRFO Lake Digi Colinvaux, P. Colinvaux et al. (1988), Bush et al. (1990) Ayauch Ecuador WTRF Lake 4 2C - Digi Bush, M. Bush et al. (1990), Bush and Colinvaux (1988), Colinvaux et al. (1988) Kumpack Ecuador Modern WTRF Lake Digi Bush, M. Bush et al. (1990) Puyo Bog Ecuador Modern WTFO Lake Digi Bush, M. Bush et al. (1990) Lake Surucucho Ecuador WTRF Lake 9 4C - Digi Colinvaux, P. Colinvaux et al. (1997) Mera Ecuador Modern WTRF Lake Digi Bush, M. Bush et al. (1990) Indanza Ecuador Modern CTRF Lake Digi Bush, M. Bush et al. (1990) Yaguara cocha Ecuador Modern CTRF Lake Raw Bush, M. Bush et al. (1990) Rum Tum Ecuador Modern CTRF Lake Digi Bush, M. Bush et al. (1990) Yambo Ecuador Modern CTRF Lake Digi Bush, M. Bush et al. (1990) Cunro Ecuador Modern CTRF Lake Digi Bush, M. Bush et al. (1990) Llaviucu Ecuador Modern CTRF Lake Digi Bush, M. Bush et al. (1990) San Marcos Ecuador Modern CGSH Mire Digi Bush, M. Bush et al. (1990) Cayambe Ecuador CGSH Mire 6 4D - Raw Graf, K. Graf, 1989 (1992) Lago Quexil Guatemala TSFO Lake 4 7D - Raw Leyden, B. W. Leyden (1984), Leyden et al. (1993, 1994) Lake Peten-Itza Guatemala TDFO Lake 7 2C - Digi Islebe, G. Islebe et al. (1996) Sierra de Cuchumatanes Guatemala Modern WAMF Pollster Digi Islebe, G. Islebe and Hooghiemstra (199) Paramo de Miranda Venezuela CGSH Mire 3 4C - Raw Salgado-Labouriau, M. L. Salgado- -Labouriau, M. L. (1988, 1991) Valle Laguna Negra Venezuela CGSH Lake Raw Graf, K. Rull et al. (1987) Paramo Piedras Blancas Venezuela CGSH Mire Raw Salgado-Labouriau, M. L. Rull et al. (1987) Sierra de Cuchumatanes 4 Guatemala Modern CTRF Pollster Digi Islebe, G. Islebe and Hooghiemstra (199) Sierra de Cuchumatanes 3 Guatemala Modern CTRF Pollster Digi Islebe, G. Islebe and Hooghiemstra (199) Sierra de Cuchumatanes 2 Guatemala Modern WAMF Pollster Digi Islebe, G. Islebe and Hooghiemstra (199) Sierra de Cuchumatanes 1 Guatemala Modern CGSH Pollster Digi Islebe, G. Islebe and Hooghiemstra (199) San Jose Chulchaca Mexico TDFO Lake 8 2C - Raw Leyden, B. W. Leyden et al. (199) Lake Coba Mexico WAMF Playa 8 7D 2C Raw Leyden, B. W. Leyden et al. (1998) Lago Catemaco Mexico TRFO Lake 2C - Raw Byrne, A. R. Byrne and Horn (1989) Lake Pátzcuaro Mexico WAMF Lake 24 2C 6C Raw Watts, W. A. Saporito (197), Watts and Bradbuy (1982) 437 Table 3. Continued. Site Country Long. Latitude Alt. Age Present Sample RC DC at DC at Data Analyst Site range biome type type publications yr BP yr BP Chalco Lake Mexico WAMF Lake 8 3C 1C Raw Lozano-Garcia, M. S. Lozano-Garcia and Ortega-Guerrero (1994), Lozano-Garcia et al. (1993), Ortega-Guerrero (1992) Lake Texcoco Mexico WAMF Lake 7 3C 4C Digi Lozano-García, S. Lozano-García and Ortega-Guerrero (in press ) Quila Mexico WAMF Lake 4 3C - Raw Almeida, L. Almeida (1997) Zempoala Mexico WAMF Lake - - Raw Almeida, L. Almeida (1997) Soberania Panama Modern TSFO Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Ocelot Pond Panama Modern TSFO Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Pipeline Rd Panama Modern TSFO Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Barro Colorado Island Panama Modern TSFO Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Panama Panama TSRO Lake 8 1C - Raw Leyden, B. W. Leyden et al. (199) El Valle Panama TSFO Lake - 2C Digi Bush, M. Bush (199), Bush and Rivera (1998), Piperno et al. (1991a, b) Cana Swamp Panama TRFO Swamp - - Digi Bush, M. Bush and Colinvaux (1994) Wodehouse Swamp Panama TRFO Swamp Digi Bush, M. Bush and Colinvaux (1994) La Yeguada, Panama TSFO Lake 11 1C - Digi Bush, M. Bush (199), Bush et al. (1992), Bush and Rivera (1998), Piperno et al. (1991a, b) Cerro Campana Panama Modern WTFO Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Cana, Darien Panama Modern TSFO Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Quebrada Nelson Panama Modern WTRF Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Horsefly Ridge Panama Modern WTRF Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Laguna Volcán Panama WTRF Lake Digi Behling, H. Behling (00) Finca Lerida Panama Modern WTRF Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Volcan Irazu Panama Modern CTRF Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Volcan Baru Panama Modern CTRF Pollster Raw Bush, M. Bush (199), Bush and Rivera (1998) Laguna Paca Peru CGSH Lake 1 4D - Raw Hansen, B. C. S. Hansen and Rodbell (199) Laguna Junin 2 Peru CGSH Lake 11 6C 1C Raw Hansen, B. C. S. Hansen and Rodbell (199) Laguna Tuctua Peru CGSH Lake 4 2C - Raw Hansen, B. C. S. Hansen et al. (1994) Laguna Milloc Peru CGSH Lake 1 3C - Raw Graf, K. Graf (1992) Laguna Pomacocha Peru CGSH Lake 4 1C - Raw Hansen, B. C. S. Hansen et al. (1994) Laguna Jeronimo Peru CGSH Lake 4 4C - Raw Hansen, B. C. S. Hansen et al. (1994) Laguna Huatacocha Peru CGSH Lake 2C - Raw Hansen, B. C. S. Hansen and Rodbell (199) Lake Valencia Venezuela STEP Lake 28 1C - Raw Leyden, B. W. Bradbury et al. (1981), Valle Leyden (198) Laguna Victoria Venezuela CGSH Lake 4 4C - Raw Graf, K. Rull et al. (1987) 438

37 Table 4. Assignment of Latin American plant functional types to Biomes. Codes Plant functional types TRFO man, tx, Te 1, Te 2, tf TSFO tx, Tr 1, Tr 2, Te 2, tf TDFO Tr 2, tf, txts, df WTRF tx, Tr 1, Te 1, wtc, ctc 2, tef, wte WEFO tx, Tr 2, wtc, ctc 2, ec, tef, wte CTRF tx, h, ctc 1, tef, wte, wte 1 WAMF Tr 2, wtc, tef, wte, ts COMI ctc 1, tef, wte 1, wte 4, ts 1 STEP sf DESE ds, df CGSH af, aa, wte 4, h CGSS af, aa, cp 439 Table. Latin American pollen taxa used in the biomisation analysis and their assignment to the PFTs. PFT codes g man tx Tr 1 Tr 2 Te 1 Pollen taxa Poaceae Acrostichum-type, Avicennia, Laguncularia, Rhizophora Alsophila, Alstroemeria, Cnemidaria, Cyathea, Dicksonia, Nephelea Alibertia, Anacardiaceae, Andira-type, Astronium, Bauhinia, Bombacaceae, Bougainvillea, Caesalpineae, Casearia-type, Clematis, Coccoloba, Copaifera, Didymopanax, Eugenia, Euterpe, Goupia-type, Guapira, Heliocarpus, Hura, Hieronima, Hippocrateaceae, Ipomoea, Laplacea, Lecythidaceae, Leguminoseae, Loranthaceae, Macharium, Macrolobium, Malpighiaceae, Malvaceae, Maytenus, Ocotea-type, Pera, Phyllostylon, Piper, Pisonia, Psychotria, Rubiaceae, Rutaceae, Salix, Sapium, Simira, Siparuna, Spirotheca, Spondias, Symplocos, Tecoma, Trema, Xylosma, Zanthoxylum Acacia, Alchornea, Alibertia, Andira-type, Anacardiaceae, Apocynaceae, Astronium, Banisteriopsis, Bauhinia, Bombacaceae, Bougainvillea, Brosimum, Brunellia, Bulnesia, Bumelia-type, Bursera, Byrsonima, Caesalpineae, Celastraceae, Chrysophyllum, Clematis, Combretaceae, Copaifera, Cordia, Coriaria, Cuphea, Curatella, Didymopanax, Elaeocarpaceae, Eryngium, Erythrina, Euterpe, Gallesia, Guapira, Hieronima, Hura, Hymenophylleace, Ipomoea, Loranthaceae, Malvaceae, Maytenus, Melastomataceae, Meliaceae, Mimosa, Passiflora, Pera, Phyllostylon, Piper, Pisonia, Protium, Pseudobombax, Rhamnaceae, Rutaceae, Salix, Sapium, Schinus, Simira, Siparuna, Spirotheca, Spondias,Styrax, Symplocos, Tiliaceae, Trema, Xylosma, Zanthoxylum Abutilon, Actinostemon concolor, Alchornea, Amanoa, Apeiba, Apocynaceae, Araliaceae, Arecaceae, Arrabidaea, Aspidosperma, Astrocarium, Begoniaceae, Bignoniaceae, Bombacaceae, Bonamia, Brosimum, Brunellia, Brownea, Calliandra, Campomanesia, Cardiospermum, Castilla, Cecropia, Cedrela, Celtis, Copaifera, Coprosma, Cucurbitaceae, Cunoniaceae, Dalbergia, Dioclea, Doliocarpus, Elaeagia, Euterpe, Ficus, Flacourtiaceae, Forsterania, Geonoma, Guapira, Guazuma, Guarea, Hura, Ilex, Inga, Iriartea, Iva xanthifolia-type, Lecythidaceae, Leguminoseae, Licania, Mabea, Macharium, Macrolobium, Macrocarpea, Malpighiaceae, Mauritia, Marcgraviaceae, Maripa, Marattia, Matayba, Melastomatacae, Meliaceae, Miconia, Moraceae, Myrsine, Myrtaceae, Mauritiella, Nyctaginaceae, Ochnaceae, Ocotea-type, Oenocarpus, Oreopanax, Palmae, Panopsis, Parahancornia, Passiflora, Plenckia, Protium, Pseudopanax laetevirens, Rauvolfia, Rhipsalis, Rubiaceae, Sapotaceae, Scheelea,Scleronema, Socratea, Sloanea, Solanaceae, Sophora, Strutanthus, Symphonia, Swartzia, Taperira, Ternstroemia cf. T. brasiliensis, Tetrochiduim, Tetraploa aristata, Thymelaeaceae, Tiliaceae, Trichilia, Trigonia, Vismia, Warsewiczia 440

38 Table. Continued. PFT codes Te 2 wtc ctc 2 ctc 1 txts h ds df tf tef Pollen taxa Abutilon, Acacia, Aegiphila, Alibertia, Apeiba, Apocynaceae, Aspidosperma, Boraginaceae, Bougainvillea, Brosimum, Brunellia, Calliandra, Cecropia, Cedrela, Celtis, Combretaceae, Croton, Cucurbitaceae, Dalbergia,Didymopanax, Dioclea, Forsterania, Hippocrateaceae, Humiria, Humulus, Ilex, Iriartea, Leguminoseae, Macrolobium, Mauritia, Melastomataceae, Miconia, Moraceae, Mysine, Myrtaceae, Ochnaceae, Ocotea-type, Palmae, Panopsis, Passiflora, Plenckia, Pseudopanax, Psychotria, Sapotaceae, Scleronema, Serjania, Sophora, Strutanthus, Swartzia, Taperira, Tiliaceae, Vismia, Warsewiczia Abies, Araucaria, Juniperus, Pinus Abies, Araucaria, Araucaria augustifolia, Cupressaceae, Dacrydium, Juniperus, Pilgerodendron, Podocarpus, Prumnopitys andina, Saxegothaea conspicua Austrocedrus chilensis, Cupressaceae, Dacrydium, Fitzroya cupressoides-type, Pilgerodendron, Pinus, Podocarpus, Prumnopitys andina, Saxegothaea conspicua, Taxodium Acacia, Aeschynomene, Agave, Anthurium, Aphelandra, Arrabidaea, Atamisquea, Ayenia, Bursera, Byrsonima, Byttneria, Cabomba, Cactaceae, Caryocar, Cayaponia, Cercidium, Chomelia, Chrysophyllum, Chuquiragua, Cissus, Clusia, Combretaceae, Convolvulaceae, Cordia, Cuphea, Curatella, Dodonaea, Echinodorus, Eichhornia, Evolvulus, Hippocrateaceae, Humulus, Hyptis, Ipomoea, Larrea, Lithraea, Malpighiaceae, Manihot, Maprounea, Menispermaceae, Metopium, Miconia, Mimosa cf. M. taimbensis, Palicourea, Pepermmia, Phaseolus, Phyllanthus, Polygala, Polylepis-Acaena, Pouteria, Portulaccaceae undiff., Prosopis, Rhamnaceae, Sapium, Schefflera, Schinus, Sebastiana, Serjania, Solanaceae, Sloanea, Stryphnodendron, Tecoma, Trixis, Zornia Arenaria, Aristotelia, Asteraceae, Berberidaceae, Berberis, Empetrum, Ericaceae, Sisyrinchium-type Ericaceae, Sisyrinchium-type Agave, Atamisquea, Cactaceae, Ephedra, Monttea aphylla Alternanthera, Ephedra, Monttea aphylla, Xyris Acalypha, Acanthaceae, Alcemilla, Alismataceae, Alsophila, Anemia, Antheroceros, Armeria, Artemisia, Assulina, Asteraceae, Astelia, Begonia, Bernardia, Brassicaceae, Bravaisia, Bromeliaceae, Calyceraceae, Caperonia, Caryophyllaceae, Cassia, Cichoriaceae, Cirsium, Cruciferae, Eriogonum, Eriocaulaceae, Euphorbia, Euphorbiaceae, Fabaceae, Geraniaceae, Gomphorena, Gunnera, Hebenaria, Iresine, Justicia, Lamiaceae, Laportea, Liquidambar, Liliaceae, Lobelia, Menispermaceae, Muehlenbeckia, Nertea, Onagraceae, Orchidaceae, Pilea, Polygala, Rhaphithamnus, Rhus, Rubiaceae, Smilax, Triumfetta, Umbelliferae, Urticaceae, Verbena, Verbenaceae, Vernonia, Viburnum, Vitis Acalypha, Acanthaceae, Apiaceae, Apium, Artemisia, Astelia, Azara, Borreria, Brassicaceae, Bravaisia, Bromeliaceae, Cassia, Cichoriaceae, Eriocaulaceae, Eriogonum, Euphorbia, Euphorbiaceae, Fabaceae, Genipa, Gordonia, Gunnera, Hippeastrum, Hydrocotyle, Iridaceae, Jungia, Justicia, Lachemella, 441 Table. Continued. PFT codes sf af cp wte wte 1 Pollen taxa Lamanonia, Lamiaceae, Laportea, Liliaceae, Lupinus, Malvaceae, Marcgraviaceae, Moritzia-type, Mutisia, Nertea, Onagraceae, Orchidaceae, Pamphalea, Perezia, Phaseolus, Pilea, Piscidia, Plantago, Polemoniaceae, Polygala, Portulaccaceae undiff., Pouteria, Ranunculaceae, Rubiaceae, Rumex, Satureja, Scrophulariaceae, Selaginella, Thalictrum, Triumfetta, Umbelliferae, Urticaceae, Verbenaceae, Vernonia, Vicia, Vitis, Wendtia, Xyris Alternanthera, Amaranthaceae/Chenopodiaceae, Antheroceros, Armeria, Assulina, Asteraceae, Astelia, Borreria, Calyceraceae, Cardus, Caryophyllaceae, Cardus, Connarus, Cruciferae, Embothrium, Eriogonum, Eryngium, Euphorbia, Euphorbiaceae, Fabaceae, Geraniaceae, Gomphorena, Gunnera, Hebenaria, Hippeastrum, Hydrocotyle, Iridaceae, Jungia, Justicia, Liliaceae, Lamiaceae, Liquidambar, Mutisia, Nanodea, Nassauvia-type, Orchidaceae, Oxalis, Phacelia, Physalis, Plantago, Polemoniaceae, Pouteria, Ranunculaceae, Restionaceae, Rosaceae, Rubiaceae, Satureja, Scutellaria-type, Umbelliferae, Urticaceae, Vicia, Vitis, Wendtia, Xyris Arenaria, Astragalus, Azorella, Bartsia-type, Borreria, Bromeliaceae, Bravaisia, Campanulaceae, Cardus, Caryophyllaceae, Deschampsia antarctica, Diphasiastrum complanatum-type, Donatia, Draba, Epilobium, Eriocaulaceae, Eriocaulon, Eriogonum, Gaimardia, Gilia, Halenia, Hebenaria, Hippeastrum, Hydrocotyle, Iridaceae, Jamesonia, Labiatae, Lachemella, Lamiaceae, Liquidambar, Lupinus, Lysipomia, Montia, Moritzia-type, Muehlenbeckia, Nassauvia-type, Orchidaceae, Oxalis, Perezia, Plantago, Puya, Quinchamalium, Relbunium, Rosaceae, Rubiaceae, Rumex, Satureja, Scrophulariaceae, Scutellaria-type, Selaginella, Sisyrinchium-type, Umbelliferae, Valeriana, Viola Apiaceae, Azorella, Gaimardia, Montia, Plantago, Saxifraga Aegiphila, Allophylus, Aphelandra, Araliaceae, Azara, Baccharis, Bauhinia, Begoniaceae, Buddleja, Bumelia-type, Clusia, Croton, Daphnopsis, Desfontainia, Elaeocarpaceae, Embothrium, Eucryphia/Caldcluvia paniculata, Euterpe, Fuchsia, Geonoma, Geraniaceae, Griselinia, Guettardia, Gunnera, Guttiferae, Hedyosmum, Heliocarpus, Humiria, Labiatae, Lomatia/Gevuina, Loranthaceae, Ludwigia, Luehea, Malpighiaceae, Matayba, Melastomatacae, Meliacea, Mimosa, Mimosa cf. M. scabrella, Mutisia, Myrica, Mysine, Nothofagus obliqua-type, Oreopanax, Palicourea, Proteaceae, Prunus, Pseudopanax laetevirens, Psychotria, Quercus, Roupala, Sambucus, Solanaceae, Stryphnodendron, Styloceras, Tepualia stipularis, Tetrochiduim, Thymelaeaceae, Trichilia, Verbenaceae, Viburnum, Warsewiczia, Weinmannia Aegiphila, Alnus, Arecaceae, Aragoa, Arcytophyllum, Aristotelia, Azara, Banara, Banisteriopsis, Begoniaceae, Bocconia, Brunellia, Buddleja, Calandrinia, Campanulaceae, Celastraceae, Chuquiraga, Clethra, Daphnopsis, Desfontainia, Dodonaea, Drimys, Epacridaceae, Ericaceae, Fuchsia, Galium, Gaultheria ulei, Geraniaceae, Gesneriaceae, Hedyosmum, Hydrangea, Hypericum, Labiatae, Loranthaceae, Ludwigia, Maytenus, Meliaceae, Meliosma, Muehlenbeckia, Myrtaceae, Mysine, Nothofagus, 442

39 Table. Continued. PFT codes wte 4 ts ts 1 aa Pollen taxa Nothofagus antarctica-type, Ostrya-type, Proteaceae, Prunus, Pseudopanax laetevirens, Quercus, Ribes, Roupala, Sambucus, Solanaceae, Tepualia stipularis, Verbenaceae, Viburnum, Weinmannia Abatia, Adesmia, Alcemilla, Alfaroa, Arcytophyllum, Assulina, Asteraceae, Clethra, Colignonia, Dodonaea, Ericaceae, Gaiadendron, Gaultheria ulei, Guttifera, Laurelia, Muehlenbeckia, Myrteola, Polylepis-Acaena, Ribes, Rosaceae, Tetrochiduim, Weinmannia Alnus, Banksia, Carpinus, Cayaponia, Fagus, Fraxinus, Juglans, Loranthaceae, Luehea, Myrica, Populus, Styrax, Trema, Ulmaceae, Vallea. Escallonia, Eugenia, Gordonia, Liquidambar, Luehea, Misodendrum, Myzodendron, Styrax, Ternstroemia cf. T. brasiliensis, Trema, Vallea Aragoa, Arcytophyllum, Arenaria, Asteraceae, Baccharis, Cruciferae, Draba, Empetrum, Ephedra, Ericaceae, Eriogonum, Escallonia, Gentiana, Gentianaceae, Hypericum, Nassauvia-type, Puya, Rosaceae, Senecio 443 Table 6. Site locations showing biome changes from the present, 6000±00 and ±00 14 C yr BP. Site Long. Latitude Alt. Present biome Modern Salina Anzotegui STEP STEP STEP Boca de Lopez TRFO TRFO Carara Biological Reserve TSFO TSFO Jotaordo TRFO TRFO Lago Crispim TRFO STEP TSFO ODP site TRFO WTRF WTRF TSFO Patagonia CTRF CTRF Reserva Volta Velha WTRF WTRF Santa Rosa TRFO TRFO Santa Rosa TSFO TSFO Tortuguero TSFO WTRF San Jose Chulchaca TDFO TDFO WEFO La Misión STEP STEP STEP Lagoa da Caço TDFO TDFO STEP Puerto del Hambre CTRF CTRF CTRF Poço Grande WTRF WTRF Harberton STEP STEP STEP Ocelot Pond TSFO WTRF Origone STEP STEP Patagonia CTRF CTRF Patagonia CTRF CTRF Patagonia CTRF CTRF Pedro Luro STEP STEP Ruta STEP STEP Ruta STEP STEP Soberania TSFO WTRF Carara WTRF WTRF Lagoa da Curuça TRFO TRFO TRFO Lake Åsa CGSH CGSH Cantarrana Swamp TSFO WTRF 444

40 Table 6. Continued. Site Long. Latitude Alt. Present biome Modern Pipeline Rd TSFO WTRF Sendro Sedro Swamp TSFO WTRF Barro Colorado Island TSFO WTRF Dichan CTRF CTRF CTRF El Camito TRFO TRFO Espuma TDFO TDFO Laguna Lincoln CTRF CTRF CTRF CGSH Laguna Lofel CTRF CTRF CTRF CGSH Laguna Six Minutes CTRF CTRF CTRF CTRF Laguna Stibnite CTRF CTRF CGSH Laguna Stibnite CTRF CTRF CTRF CGSH Patagonia CTRF CTRF Patagonia CTRF CTRF Patagonia CTRF CTRF Patagonia CTRF CTRF Patagonia CTRF CTRF Puerto Eden CTRF CTRF CTRF Rio (unclear) TDFO CTRF Rio Jaguaribe I TDFO STEP Rio Jaguaribe II TDFO TDFO Estero Huitanque CTRF CTRF COMI Laguna Palmita TSFO WTRF Patagonia CTRF CTRF Patagonia CTRF CTRF Campina Grande I TSFO STEP Mirim TDFO STEP Patagonia CTRF CTRF Picos TDFO WTRF Rio Mirim TDFO WTRF Lagoa Grande TDFO CTRF Mayol CTRF CTRF CTRF Punta Arenas CTRF CTRF CTRF CGSH Rano Raraku Bore TDFO TDFO TDFO TDFO 44 Table 6. Continued. Site Long. Latitude Alt. Present biome Modern La Selva, Heredia TSFO TSFO Patagonia CTRF CTRF Rio de Contas TDFO STEP Rio Jequitinhonha TDFO TDFO Rio Protengi TDFO CTRF Rio São Francisco TDFO STEP Arroyo Sauce Chico STEP STEP Gaviotas TDFO TDFO Lago Bolim TDFO STEP Lake Coba WAMF WAMF WEFO Panama TSFO WTRF TSFO Patagonia CTRF CTRF Patagonia CTRF CTRF Torres del Paine CTRF CTRF TDFO Empalme Querandíes STEP STEP TDFO La Pacifica, Guanacaste TSFO TSFO Lago Quexil TSFO WAMF WEFO WAMF Puchilco CTRF CTRF CTRF Rano Kao TDFO TDFO Ruta STEP STEP Puerto Octay PM CTRF CTRF CTRF COMI Comprida WTRF WTRF TRFO Geral WTRF WTRF TRFO Chepu CTRF CTRF Carajas TSFO TSFO TSFO TSFO Patagonia CTRF CTRF Patagonia CTRF CTRF Mariñame-II TRFO TRFO WTRF Monica TRFO TRFO TRFO Carimagua TDFO TDFO CGSS Patagonia CTRF CTRF Patagonia CTRF CTRF Patagonia CTRF CTRF Patagonia CTRF CTRF 446

41 Table 6. Continued. Site Long. Latitude Alt. Present biome Modern Patagonia CTRF CTRF Patagonia CTRF CTRF Sardinas TDFO TDFO TDFO El Piñal TDFO TDFO TDFO TDFO Patagonia CTRF CTRF Patagonia CTRF CTRF Patagonia CTRF CTRF Patagonia CTRF CTRF Patagonia CTRF CTRF Aguads Emendadas TDFO CTRF TSFO CGSH Atlantic WTRF WTRF Cerro La China STEP TDFO TDFO Crominia TDFO TDFO TSFO TDFO Lake Peten-Itza TDFO TDFO WEFO Moreno Glacier Bog CTRF CTRF COMI Patagonia CTRF CTRF Piusbi TRFO TRFO TRFO Laguna Angel TDFO TSFO TDFO LagoBsAs CTRF WAMF Limoncocha TRFO CTRF Lago Agua Sucia TDFO TDFO STEP Cataracta, Carara TSFO WTRF Cataracta, Carara TSFO WTRF Añangucocha TRFO WTRF Cuyabeno TRFO WTRF Escondido TSFO WTRF Santa Rosa TSFO TSFO Santa Rosa TSFO WTRF Cafetal, Guanacaste TSFO WTRF La Pata TRFO TSFO WTRF WTRF Lagoa das Patas TRFO WTRF TSFO WTRF Loma Linda TDFO TDFO TDFO La Esperanza CTRF CTRF 447 Table 6. Continued. Site Long. Latitude Alt. Present biome Modern Lago Agrio TRFO WTRF Lake Agrio TRFO TRFO Lake Santa Cecilia TRFO TRFO Santa Cecilia TRFO WTRF Lago Catemaco TRFO WAMF Cuiaba TDFO TDFO Laguna Bonilla TSFO WTRF Laguna Río Cuarto WTRF WTRF Lago do Pires TSFO TSFO TSFO Rio São Francisco TDFO TDFO TSFO Lake Valencia STEP STEP TDFO Rano Aroui TDFO TDFO CGSH WTRF Saquinho TSFO TSFO WEFO Cana Swamp TRFO TRFO Caunahue CTRF CTRF El Valle TSFO TSFO TDFO Wodehouse Swamp TRFO TRFO Assis TSFO TSFO Ayauch WTRF WTRF TSFO Bauru TSFO TSFO Laguna La Palma WTRF WTRF Laguna Cedeño WTRF WTRF Brauillo Carillo, Heredia WTRF WTRF Lagoa Santa TDFO TDFO Pico Salam STEP STEP La Yeguada TSFO TSFO TSFO San Pedro CTRF CTRF Botucatu WTRF WTRF Brotas WTRF WTRF Curcuab WTRF WTRF Kumpack WTRF WTRF Laguna González WTRF WTRF Bosque Alegre WTRF WTRF 448

42 Table 6. Continued. Site Long. Latitude Alt. Present biome Modern Laguna Congo WTRF WTRF Laguna Hule WTRF WTRF Lagoa Grande TDFO CTRF Katira TDFO TDFO TDFO TDFO Laguna Bella Vista TSFO TSFO TSFO TDFO Laguna Chaplin TSFO TSFO TDFO TSFO Laguna María Aguilar WTRF WTRF AlercesNor CTRF CGSH Cerro Campana WTRF WTRF Mallin Book CTRF COMI CTRF Primavera CTRF CGSH TDFO Rio da Curuá WTRF TRFO Comallo CGSH STEP Colombo TSFO TSFO Puyo Bog WTRF WTRF Encantado CTRF COMI Lake Surucucho WTRF CTRF WTRF Meseta Latorre CGSH COMI CTRF Cana, Darien TSFO TSFO Meseta Latorre CGSH COMI CGSH Volcán Cacao WTRF WTRF Brasilia TDFO CTRF Cueva Haichol STEP STEP STEP Salitre WTRF WTRF TDFO CGSH AustroEsqu CTRF CTRF Mera WTRF WTRF Quebrada Nelson WTRF WTRF Horsefly Ridge WTRF WTRF Serra da Boa Vista WTRF WTRF CTRF Serra Campos Gerais WTRF WAMF TDFO Pitalito WEFO WEFO TSFO Serra do Rio Rastro WTRF WTRF CGSH Vaca Lauquen CTRF COMI CGSH 449 Table 6. Continued. Site Long. Latitude Alt. Present biome Modern Vaca Lauquen CTRF COMI CGSH Laguna Volcán WTRF WAMF Monteverde, Heredia WTRF WTRF Finca Lerida WTRF WAMF Piagua WEFO WEFO WEFO Pantano de Genagra WEFO TDFO WEFO Rio Timbio WEFO WEFO WEFO Morro da Igreja WTRF WTRF WTRF Libano WEFO WEFO COMI Morro de Itapeva WTRF WEFO TDFO CGSH Veranada Pelan CGSH CGSH TDFO de Pedro Palo III COMI WTRF WTRF Herrera COMI CGSH COMI CTRF Salina CGSH STEP STEP Ubaqué WEFO WEFO Lake Pátzcuaro WAMF WAMF WAMF WAMF Indanza CTRF WTRF Yaguara cocha CTRF WTRF Chalco Lake WAMF WAMF WAMF WAMF Volcan Irazu CTRF CTRF La Chonta CTRF CTRF COMI WAMF Lake Texcoco WAMF WAMF WAMF WAMF Rum Tum CTRF WTRF Talamancas CTRF WAMF Ciudad Universitaria X COMI COMI WAMF WAMF El Abra II COMI CTRF CTRF Fúquene II COMI CGSH CTRF CTRF Volcan Poas CTRF CTRF Laguna Botos CTRF WTRF Volcan Baru CTRF CTRF Yambo CTRF WTRF Bog CTRF WTRF Bog CTRF WTRF 40

43 Table 6. Continued. Site Long. Latitude Alt. Present biome Modern Tres de Junio CTRF CTRF Wasa Mayu COMI CGSH STEP Cunro CTRF WTRF Quila WAMF WAMF WAMF Sierra de Cuchumatanes WAMF WAMF Veranada Vulkanpickel CGSH STEP TDFO Laguna Barva CTRF WTRF Sierra de Cuchumatanes CTRF CTRF Quebrador CTRF CTRF Alsacia COMI WAMF COMI Zempoala WAMF WAMF Llaviucu CTRF WTRF Salado CGSH CGSH Agua Blanca COMI COMI CTRF CGSH Valle Laguna Victoria CGSH CGSH CGSH Paramo de Miranda CGSH CGSH CGSH Cienaga del Visitador COMI COMI CGSH Asuncion CTRF CTRF San Marcos CGSH TDFO Sierra de Cuchumatanes CTRF CTRF La Guitarra COMI COMI CTRF Valle Laguna Negra CGSH CGSH Lago de las Morrenas CTRF WAMF CTRF Ciega I COMI CTRF Lago Chirripó CTRF WAMF La Primavera CGSH CGSH COMI De la América CGSH CGSH CTRF Paramo Palacio CGSH CGSH CGSH Andabobos CGSH CGSH CGSH Laguna Paca CGSH CTRF Sierra de Cuchumatanes WAMF WAMF Paramo de Laguna Verde CGSH CGSH CGSH 41 Table 6. Continued. Site Long. Latitude Alt. Present biome Modern Paramo de Peña Negra CGSH CGSH COMI El Bosque CGSH CGSH Turbera de Calostros CGSH CGSH Lake Huinãmimarca CGSH CGSH CGSH CGSS Bobos CGSH CGSH El Gobernador CTRF CTRF WTRF Tumbre CGSH CGSH STEP Valle de Lagunillas CGSH CGSH CTRF Cerro Calvario CGSH CGSH CGSH Aguilar CGSH CTRF STEP Amarete CGSH CGSH CGSH Greja CGSH CGSH CGSH La Rabona CGSH CGSH CTRF Rio Kaluyo CGSH CGSH STEP Paramo Piedras Blancas CGSH CGSH CGSH Corazón Partido CGSH CGSH El Trinagulo CGSH CGSH Laguna Junin CGSH COMI COMI CGSH Sierra de Cuchumatanes CGSH CTRF Aguas Calientas CGSH CGSH CGSH Laguna Tuctua CGSH COMI WTRF Sajama CGSH CGSH Laguna Milloc CGSH CGSH CGSH Cayambe CGSH CGSH CGSH Cotapampa CGSH CGSH CGSH Laguna Jeronimo CGSH CGSH CGSH Laguna Pomacocha CGSH CGSH CGSH Laguna Huatacocha CGSH CGSH CGSH Cumre Unduavi CGSS CGSS CGSS Ajata CGSH CGSH Chacaltaya CGSH CGSH CGSH Mt. Blanco CGSS CGSS Katantica CGSH CGSH CGSH 42

44 Figure 1. Map of Latin America depicting the present-day summer and winter position of the ITCZ and the macroscale wind (and hence moisture) patterns over Latin America ITCZ position July ITCZ position January Prevailing wind direction 0 40 Fig. 1. Map of Latin America depicting the present-day summer and winter position of the ITCZ and the macroscale wind (and hence moisture) patterns over Latin America Figure 2. Map of the modern potential vegetation as derived from Schmithüsen, J. (1976) Hück (1960). For example, the various divisions of seasonally dry forest such as Cerradõ, Caatinga, Campo Rupstre, Savanna, are combined to the biome of tropical dry forest. Fig. 2. Map of the modern potential vegetation as derived from Schmithüsen (1976) and Hück (1960). For example, the various divisions of seasonally dry forest such as Cerradõ, Caatinga, Campo Rupstre, Savanna, are combined to the biome of tropical dry forest

45 forest (j, k), dominated by mangrove (l), tropical seasonal forest (m), cool temperate forest (j, k). Plate 0 shows the multi-vegetated lays within cloud forest (cool temperate rainforest), some of these taxa such as Rhipsalis and Bromeliacae are also found in very dry ecosystems. The bottom plate (l) shows the importance of edpahic factors on controlling vegetation; in this case local hydrology where rill channels allow trees to grow in areas that would be dominated by grassland. c a b e g i f j m d h l k o n p J Fig. 3. Examples of biomes used in Latin America from cool grass shrubland from the paramo of Colombia (a, b) to cool mixed forest (c, d, e, f), tropical dry forest (g, h) with dominance of steppe (i) tropical rain forest (j, k), dominated by mangrove (l), tropical seasonal forest (m), cool temperate forest (j, k). Plate 0 shows the multi-vegetated lays within cloud forest (cool temperate rainforest), some of these taxa such as Rhipsalis and Bromeliacae are also found in very dry ecosystems. The bottom plate (l) shows the importance of edpahic factors on controlling vegetation; in this case local hydrology where rill channels allow trees to grow in areas that would be dominated by grassland Figure 4. Cross an altitudinal cross section of the Andes showing the standard vegetation units and their relationship to Biomes (a) and plant functional types (b). CGSS 000 (a) CGSH 4000 Altitude (m) COMI 3000 CEFO WAMF 00 WEFO 00 TRFO TSFO TDFO STEP 0 TROPICAL RAIN FOREST UPPER ANDEAN FOREST SAVANNA SUB-PARAMO LOWER ANDEAN FOREST PARAMO / SUPER PARAMO 000 cp (b) Altitude (m) wte4 af 4000 aa ts ctc wte1 g ctc1 ec 00 wte tf tx tf 00 Tr1 Te1 sf Te2 Tr2 txts man 0 Fig. 4. Cross an altitudinal cross section of the Andes showing the standard vegetation units and their relationship to Biomes (a) and plant functional types (b). 46 1